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ulletin of the 



British 




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1^ 



Ornithologists' 

Club 




MEETINGS are nomially held in the Sherfield Building of Imperial College, South Kensington, London 
SW7. The nearest Tube station is at South Kensington; a map of the area will be sent to members, on request. 
(Limited car parking facilities can be reserved [at a special reduced charge of £5.00], on prior application to 
the Hon. Secretary.) 

The cash bar is open from 6.15 pm, and a buffet supper, of two courses followed by coffee, is served at 
7.00 pm. (A vegetarian menu can be arranged if ordered at the time of booking.) Infonnal talks are given on 
completion, commencing at about 8.00 pm. 

Dinner charges are £21.00 per person as from 1 January 2006. 



FORTHCOMING MEETINGS 

See also BOG website: http://www.boc-online.org 



26 September 2006 — Please note that the proposed talk for this date (Lars Svensson — The Almaty skin col- 
lection and the birds of Kazakhstan) has been cancelled. An alternative speaker or programme of short talks 
will be substituted as soon as possible. It is hoped that the revised programme can be announced at the June 
meeting and also be published in the September issue of the Bulletin scheduled for dispatch on 12 
September. Under these circumstances, notice to attend the September meeting will be accepted up to one 
week in advance (see below). 

Applications to Hon. Secretary (address below) by 17 September please 



7 November 2006 — Edward C. Dickinson — Avian nomenclature and the ICZN Code: a layman 's view. 
Edward Dickinson is probably best known as the editor of The Howard & Moore complete checklist of birds 
of the world, third edition (2003). The talk will examine some of the ambiguities and difficulties that affect 
decisions on authorship and on the dating of bird names, and open up discussion of changes that are or may 
be under consideration. 



Applications to Hon. Secretary (address below) by 24 October 



5 December 2006 — Prof. Jeremy Greenwood — The future of birds and Man. Jeremy Greenwood has been 
Director of the British Trust for Ornithology since 1988. He has worked on genetics, behaviour, ecology and 
biogeography of snails and various birds, particularly Guillemots Uria aalge. Snow Buntings Plectrophenax 
nivalis and Blue Tits Parus caeruleus. Starting from an ornithological base, Jeremy will consider the 
prospects facing mankind and the environment from the viewpoint of ecological science. 

Applications to Hon. Secretary (address below) by 14 November 



Overseas Members visiting Britain are especially welcome at these meetings, and the Hon. Secretary would 
be very pleased to hear from anyone who can offer to talk to the Club giving as much advance notice as 
possible — please contact: S. A. H. (Tony) Statham, Ashlyns Lodge, Chesham Road, Berkhamsted, Herts. 
HP4 2ST, UK. Tel. 01442 876995 (or e-mail: boc.sec@bou.org.uk). 



Club Announcements 81 Bull. B.O.C. 2006 126(2) 

Bulletin of the 

BRITISH ORNITHOLOGISTS' CLUB 

Vol. 1 26 No. 2 Published 5 June 2006 

CLUB ANNOUNCEMENTS 

Members are reminded that subscriptions for 2006 fell due on 1 January. Details for membership 
correspondence, subscriptions and applications can be found on the inside back cover. All such issues 
should now be addressed to the BOC Office, P.O. Box 417, Peterborough PE7 3FX, e-mail: 
boc.office@bou.org.uk 

Correspondence on all general Club matters should continue to be addressed to: Hon, Secretary, 
S. A. H. (Tony) Statham, Ashlyns Lodge, Chesham Road, Berkhamsted, Herts. HP4 2ST, e-mail: 
boc.sec@bou.org.uk 

Congratulations to Peter Tate on achieving 50 consecutive years (1956-2006) membership, who 
therefore becomes a Hon. Life Member. We are grateful for his 12-year service as Hon. Treasurer 
(1962-74). 

Committee is pleased to welcome the following new members who were formally elected at the 
meeting held on 25 April: J. del Hoyo (Spain), J. L. Garrett (USA), M. Lentino (Venezuela), S. Peters 
(UK), B. O'Brien (UK) and Dr R. Winkler (Switzerland) 

The 935th meeting of the Club was held on Tuesday 24 January 2006, in the Sherfield Building Annexe, 
Imperial College, London. Seventeen members and nine guests were present. 

Members attending were: Cdr. M. B. CASEMENT, RN {Chairman), Miss H. BAKER, D. R. 
CALDER, S. E. CHAPMAN, Prof R. A. CHEKE, Dr J. COOPER, J. B. FISHER, Revd. T. W. 
GLADWIN, D. GRIFFIN, Dr J. P HUME, Dr C. F. MANN, D. J. MONTIER, R J. SELLAR, S. A. H. 
STATHAM, C. W. R. STOREY, Cdr. F. S. WARD, RN, and R J. WILKINSON 

Guests attending were: Mrs J. CALDER, Mrs C. R. CASEMENT, Mrs R A. CHAPMAN, Field 
Marshal Sir JOHN CHAPPLE {Speaker), Vice Admiral Sir DAVID DOBSON, Mrs J. M. GLADWIN, 
Mrs M. MONTIER, B. O'BRIEN and Mrs J. WARD. 

After dinner, Sir John Chappie presented his Reminiscences of a nature conservationist, a summary 
of how birds have influenced his life with examples of how he has repaid the 'debt'. His earliest experi- 
ences of being shown nests of Spotted Flycatcher Muscicapa striata and Chiffchaff Phylloscopus colly- 
bita hooked him for life. For his ninth birthday, he received the five-volume Handbook of British birds 
(Witherby et al), and after the war, aged 16, he participated in an expedition to Newfoundland. Before 
the expedition, he spent time at the Natural History Museum in South Kensington learning skinning and 
preservation techniques. Sir John was made a Fellow of the Zoological Society of London and the Royal 
Geographic Society whilst still a teenager and turned these into life memberships as soon as he could 
afford to do so. Much later, he became President of the former (1992-95) and a Council Member of the 
latter (1980-2000). His extensive military and diplomatic career also led to memberships of numerous 
ornithological and conservation-minded organisations, giving him the opportunity to see many of the 
world's rare species including the Pink-headed Duck Rhodonessa caryophyllacea at Alfred Ezra's 
Foxwarren Park in Surrey. 

Sometimes when visiting overseas military colleagues. Sir John's interest in ornithology was met 
with bewilderment. Some armies would regard this weird hobby as an automatic bar to promotion and in 
a military context such interests could be frustrating, such as being on patrol in Borneo with 1 00 soldiers 
and trying to stop the whole company for a special bird. Conversely it always seemed that one might hear 
a tantalising new species when waiting in ambush, but be unable to get close enough to identify it. 



Club Announcements 82 Bull. B.O.C. 2006 126(2) 

Sir John's tenure in Hong Kong began in the late 1950s when the New Territories were very rural. 
Old villages were distinct entities surrounded by paddyfields. Much of the environment was man-made 
and uncontrolled shooting, trapping and netting were also widespread. Despite the enormous changes in 
agriculture that developed from the 1 960s, a few people persuaded the local District Officer that Mai Po 
marsh should be made a conservation area. The border fencing to control illegal immigrants and the con- 
stant military presence to enforce law and order actually helped protect the area. The founding of the 
Hong Kong chapter of the World Wildlife Fund provided a substantial boost post- 1980 and today the 
reserve is well supported and attracts many visitors. 

Sir John concluded with an outline of the work to establish a captive-breeding programme for three 
Gyps species of vultures in Nepal. Populations have been decimated by the use of Diclofenac as a pain 
reliever for cattle. Several organisations are taking the lead on this project, including the King Mahendra 
Trust and, whilst bureaucracy has frustrated preliminary efforts, progress is being made. Diclofenac has 
now been banned in both India and Nepal, and a substitute drug (Meloxican), regarded as harmless to 
vultures, is being introduced. 

The 936th meeting of the Club was held on Tuesday 14 March 2006, in the Sherfield Building Annexe, 
Imperial College, London. Nineteen members and seven guests were present. 

Members attending were: Cdr. M. B. CASEMENT, RN (Chairman), Miss H. BAKER, Prof R. A. 
CHEKE, Ms K. COOK, Dr J. COOPER, J. B. FISHER, F. M. GAUNTLETT, A. GIBBS, D. GRIFFIN, 
R. H. KETTLE, R. R. LANGLEY, Dr C. F. MANN, D. J. MONTIER, R J. OLIVER, R. C. PRICE, Dr 
R. P PRYS-JONES, P J. SELLAR, S. A. H. STATHAM and P J. WILKINSON. 

Guests attending were: Mrs C. R. CASEMENT, Mrs B. FISHER, Mrs M. H. GAUNTLETT, Mrs M. 
MONTIER, R. PERRON {Speaker), Dr S. TURVEY and Mrs S. VOGEL. 

After dinner, Richard Perron talked on The phantoms of the rainforest — the genus Casuarius. The 
genus Casuarius comprises three extant species, which, although very similar, can readily be separated 
by the number of wattles on the foreneck, the size and shape of the casque on the head and distinctive 
colouring on the throat, shoulders and neck. Cassowaries are ratites, related to ostriches, rheas, kiwis and 
emus. They, along with the extinct elephant birds and moas, share a common flying ancestor which lived 
c. 1 00 M YA and have reached their present locations through continental drift. Cassowaries range through 
dense forests of north-east Queensland, New Guinea and several nearby large islands, including New 
Britain and Seram, where populations of C. bennetti and C casuarius, respectively, have been introduced 
by man, albeit probably hundreds of years ago. Adults are uniformly black with double feathers like their 
closest relative, emus. They are flightless, but have a remnant wing with six shafts, popularly used by 
New Guinea tribesmen as nose decoration. They are very shy, but can be very dangerous and have killed 
humans. The male alone incubates the 3-5 eggs for c.54 days, neither drinking nor eating during this peri- 
od. Young are precocial and brown with dark parallel longitudinal stripes on the back. They stay with the 
male until sexual maturity at c. 18 months, when they establish their own territories. Females are normal- 
ly larger than males and adults can reach 1.8 m tall and weigh 55 kg. Cassowaries feed on forest fruits, 
but also take insects and small animals. Aboriginal life in Queensland and New Guinea is inseparably 
intertwined with the cassowary, which forms part of religious, cultural and economic affairs. With no nat- 
ural enemies, cassowaries are threatened by deforestation, introduced species and overhunting, though 
efforts in Queensland are underway to protect them. There are c.600 cassowaries in captivity worldwide, 
almost all of them C. casuarius, but there is no realistic captive-breeding programme. 

Revd. Geoffrey K. McCulloch, QBE 

It is with great regret, we report Geoffrey's death, aged 94, on 6 November 2004. A member of the Club 
since 1944, he became a Hon. Life Member in 1994. Bom in the Wirral, he moved to Northwood, 
Middlesex, in 1931. He was called to the Bar in 1935 and shortly after the outbreak of war was 
commissioned into the Rifle Brigade. In 1943 he was seconded to the Anny Legal Services, travelling 
widely around Europe in the aftermath of the war. He was partly responsible for re-writing the Manual 
of Military Law and was awarded the OBE in 1 957. In 1962 he retired, with the rank of Colonel, to begin 



Club Announcements 83 Bull. B.O.C. 2006 126(2) 

training for the Church of England. Ordained in 1 964, he became Vicar of St Matthew with St Barnabas, 
Hull. On his retirement in 1979, he returned to Northwood, where he remained active as a Hon. Member 
of Staff at Emmanuel Church, and was a regular at Club dinners. He served on the Committee 1981-83, 
as Vice-Chairman 1983-86 and finally as Chairman 1986-89. 

Geoffrey's passion for ornithology began in the Wirral and he could fondly remember the exact spot 
he had found his very first nest! He pursued this interest throughout his period in the army. During his 
time in Egypt, in the early 1950s, he and a colleague were once looking for nests opposite the 
commander-in-chief's residence. A guardsman on sentry duty, who was obviously bored, presented arms 
every time Geoffrey appeared out of a ditch or hollow — often several times a minute! Whether abroad or 
at home, Geoffrey had his binoculars always to hand. He was married to Isabel for 55 years and they had 
two children. 

Michael Casement 



ANNUAL GENERAL MEETING 2006 

\. The Annual General Meeting of the British Ornithologists' Club was held in the Sherfield Building 
Annexe, Imperial College, London SW7, on Tuesday 25 April 2006 at 6 pm with Cdr. M. B. Casement 
OBE, RN, in the Chair. Seventeen members were present (including six from the committee) and 
apologies were received from: I. R. Bishop, R. J. Dowsett, S. R Dudley, J. B. Fisher, S. Lowe and R 
Sellar. The Minutes of the 2005 Annual General Meeting held on 26 April 2005, which had been 
published {Bull. Brit. Orn. CI. 125: 82-84), were approved and signed by the Chairman. 

2. Chairman's report. This is my first report, as your Chairman, but I propose to be very brief, because 
most of the points are already covered in the Trustees' Annual Report, and will be covered in more detail 
by David Montier, Guy Kirwan and Tom Gladwin, under the appropriate items to follow. My thanks are 
due to each of them for their key roles in managing our affairs, and also to Tony Statham who has done 
a splendid job in continuing the tasks he inherited from me as Hon. Secretary. In summary, it has been a 
busy year behind the scenes and on completion of the increased subscription renewals, as from 1 January 
2005, responsibility for all membership matters was transferred from myself to Steve Dudley in July. 
Steve Dudley also took over responsibility from Paul Salaman for the management of the website; some 
improvements have been made, including facilities for online payments, and others are planned. The 
steady and continuing small decline in membership is a matter of concern, but is similar to other 
comparable organisations. One regrettable casualty of these changes is that we can no longer produce the 
annual list of members. Members seeking contact details and address changes of other members should 
contact Steve Dudley, direct. 

As you will see in the report, our financial affairs are in a very healthy state, largely thanks to the 
expert management of our Hon. Treasurer, and also through a welcome rise in the stock market. We again 
held seven evening meetings, on a wide range of subjects, and a similar programme is planned for 2006. 
Our thanks are due to Imperial College for their excellent administrative support and, despite the annual 
rise in dinner charges, for continuing to allow us very favourable rates. The Club's projector has been put 
to good use throughout the year, and we are grateful to Pat Sellar and Ron Kettle for their technical skills 
with the sound equipment. 

The BOC-BOU Joint Publications Committee (JPC), under the chairmanship of Tom Gladwin, has 
made continuing progress, the highlight of which was the publication of the Bird Atlas of Uganda. Bob 
Cheke has filled the vacancy left by the late Janet Kear, as Commissioning Editor of the publications in 
the Checklist series. 

As Guy Kirwan will report, the Bulletin has been maintained at its usual high standard. Our thanks 
are due to Mary Muller for continuing to produce the annual index; she has now done this for more years 
than I can remember, and has asked to be relieved of this task — action is in hand to seek her replacement. 
The most important step has been, as announced in Bull Brit. Orn. CI. 126: 1, the establishment of a 
Bulletin Subcommittee, under the chairmanship of Prof Chris Feare, with terms of reference to develop 
and recommend policies for all aspects of the management of the Bulletin. High on their agenda is to 



Club Announcements 84 Bull. B.O.C. 2006 126(2) 

investigate the practicalities of producing an online version. The subcommittee is due to meet on 
Wednesday 3 May. 

Your Committee is also looking at a long-term 'Vision for the Future' and the Bulletin 
Subcommittee's recommendations will be a major contribution to this study. I shall give a brief explana- 
tion of this later this evening. This time next year, we may well have some fairly major recommendations 
and changes to place before you. 

3. Trustees Annual Report. The Chairman advised the meeting that the Trustees Annual Report and 
Annual Accounts were again combined in the handout available to those attending the meeting and 
confirmed that these would also be published in the June issue of the Bulletin. The Hon. Treasurer drew 
attention to the income and expenditure details shown on pp. 8-9 of the accounts and summarised the 
balance sheet shown on p. 7. The Chairman thanked the Hon. Treasurer for his support and seconded his 
proposal that the accounts should be formally accepted; all those present agreed. 

A brief overview was also given for the Barrington Fund whose assets stood at £1,162 as of 31 
December 2005. The Hon. Treasurer proposed that this balance should be transferred to general funds, 
which was seconded by the Chairman and agreed unanimously by those present. 

4. The Bulletin. Some 43 papers were pubHshed as part of Vol. 125, including type descriptions of two 
new taxa. The interval between receipt and publication was 6-18 months, averaging c. 12 months, a slight 
improvement on most recent years, although it is hoped to reduce 'waiting time' slightly further. During 
the final part of the year, work was progressed towards publication of the Recent Avian Extinctions 
conference proceedings, which will be published as a Supplement, timed to appear with the June 2006 
issue of the Bulletin. The Hon Editor thanked Eng-Li Green, of Alcedo Publishing, Mike Dawson and 
Crowes, Mary Muller, Tony Statham and staff at the Natural History Museum, Tring, for their assistance. 

5. Publications report. Revd. Tom Gladwin, Chairman of the Joint Publications Committee (JPC), drew 
attention to the relevant section in the Trustees Annual Report and confirmed that the publication of The 
Birds of Sao Tome and Principe with Annobon was imminent. Future publications planned for 2006 
include The Birds of Borneo and The Birds of Barbados, and both manuscripts are now with 
Commissioning Editor Prof Bob Cheke. The JPC is seeking advice in order to make recommendations 
that authors should follow an established and authorised taxonomic system or otherwise with the JPC's 
consent. Some research is also being undertaken to explore online printing and printing on demand. 

6. Election of Officers and Committee. The Chairman said that the election of Officers was as proposed 
in the agenda as published in Bull. Brit. Orn. CI. 126: 1. 

Mr S. A. H. Statham be re-elected as Hon. Secretary. 
Mr D. J. Montier be re-elected as Hon. Treasurer 

No other changes to the Committee are proposed, as all other members are eligible to serve at least 
one more year in office. 

The following ex-officio members were confirmed in continuation: 

Revd. T. W. Gladwin {Chairman Joint Publications Committee) 

Prof R. A. Cheke {Hon. Publications Officer) 

S. P. Dudley {Hon. Website Manager) 

G. M. Kirwan {Hon. Editor) 

Prof. C. J. Feare {Chairman Bulletin Subcommittee) 

7. Any Other Business. The Chairman advised that the Hon. Secretary (Tony Statham) had received a 
note from F. M. Gauntlett (FMG) expressing concern about an article in Bull. Brit. Orn. CI. 125 (3) 
(Robbins, M. B. & Zimmer, K. J. Taxonomy, vocalisations and natural history of Philydor dimidiatum 
(Fumariidae), with comments on the systematics of Syndactyla and Simoxenops). FMG stated that the 
article implied that the Club condoned the collection of specimens by the authors in a national park 
without having the relevant permit(s). The Chairman said that the Committee had already discussed this 



Club Announcements 85 Bull. B.O.C. 2006 126(2) 

matter and that the Hon. Editor (G. M. Kirwan) had corresponded with FMG. It was agreed that future 
articles should confirm that the necessary permit(s) had been obtained; it was noted that a similar policy 
was adopted for articles in Ibis. 

The meeting closed at 6.45 pm. 

BRITISH ORNITHOLOGISTS' CLUB 

Founded 5 October 1892 

Registered Charity No. 279583 

TRUSTEES' ANNUAL REPORT FOR 2005 

List of Trustees — Committee 

Cdr. M.B. CASEMENT, QBE, RN Chairman (2005) 

Miss H. BAKER Vice-Chairman (2005) 

S. A. H. STATHAM Hon. Secretary (2004) 

D. J. MONTIER Hon. Treasurer {\991) 

I. R. BISHOP, QBE (2003) 

Dr J. H COOPER (2005) 

Dr J. R HUME (2004) 

C. W. R. STOREY (2003) 

R J. WILKINSON (2005) 

Ex-officio Revd. T. W. GLADWIN Chairman JPC (2002). 

Correspondence and enquiries to the Hon. Secretary, Ashlyns Lodge, Chesham Road, Berkhamsted, 

Herts. HP4 2ST 
Auditors and Independent Examiners Porritt Rainey, 9 Pembroke Road, Sevenoaks, Kent, TNI 3 IXR 
Bankers Barclays Bank pic. Dale House, Wavertree Boulevard, Liverpool, L7 9PQ 

Constitution 

The British Ornithologists' Club was founded in October 1892. It currently operates under Rules revised 
in 2000 and approved at a Special General Meeting on 31 October of that year. Members of the 
Committee, who are also the Trustees of the Club, are listed above with the dates of their appointment. 
Committee members are elected for a term of four years, with the exception of the Hon. Secretary and 
Hon. Treasurer, who are both subject to re-election at each Annual General Meeting. 

Objects of the Charity 

The promotion of scientific discussion between Members of the British Ornithologists' Union (BOU) and 
others interested in ornithology, and to facilitate the dissemination of scientific information concerned 
with ornithology, with a particular emphasis on avian systematics, taxonomy and distribution. 

The Committee 

The Committee met six times during the year. At the AGM on 26 April Michael Casement was appointed 
Chairman in succession to Dr Clive Mann and agreed to remain Membership Secretary until this position 
was handed over to Steve Dudley, the BOU Administrator, under the terms of a joint administration 
agreement between the Club and the BOU. This took effect from 1 August but the subscription 
management of BOC Institutional Subscribers and sales of BOC publications commenced from 1 
January. Other appointments at the AGM included Helen Baker as Vice-Chairman {vice Michael 
Casement) and Dr Joanne Cooper and Peter Wilkinson in succession to Dr Paul Salaman and Prof Bob 
Cheke, who remains Hon. Publications Officer. To assist them in understanding their responsibilities as 
Trustees, all Committee members have been provided with a copy of two booklets issued by the Charity 
Commissioners outlining matters of which Trustees need to be aware. 



Club Announcements 86 Bull. B.O.C. 2006 126(2) 

Dr Paul Salaman, Hon. Website Manager, continued to maintain the BOC website until August when 
this responsibility was also handed over to Steve Dudley under the joint administration agreement with 
the BOU. Steve Gregory, Paul Salaman and Alistair Carr continued to progress the project for the elec- 
tronic archiving and marketing of the complete set of the Bulletin. 

The Committee continues to be most grateful to the Trustees of the Herbert Stevens Trust Fund 
(Nigel Crocker, Peter Oliver and Richard Price) for their time and expertise advising on the performance 
of the Fund. 

Meetings 

The number of evening meetings held at Imperial College, London, was maintained at seven in 2005. A 
total of 196 (151 members and 45 guests) attended these meetings, which represented an average 
attendance of 28. The programme of speakers during the year again covered a wide variety of 
ornithological subjects, including the songs of North American sparrows and warblers, the use of shade 
by birds in hot climates, a collection of mummified birds in the Natural History Museum, the spread of 
the House Crow Corvus splendens, ship-based observations of seabird and landbird migration in the 
eastern Atlantic, and the parrots of the Mascarenes. As in previous years, the 26 April meeting following 
the AGM took the form of a social evening, during which informal short talks and brief discussions were 
contributed by nine participants on a range of subjects from individual species to Lord Rothschild's 1905 
Red List of extinct and endangered species, and an overview of the Royal Naval Birdwatching Society 
database. Summaries of these talks are to be found in the Club Announcements section of the Bulletin. 
Unfortunately, Imperial College increased their catering and venue charges again for 2005 requiring an 
increase in the cost of Club dinners to £20 per head. 

The Bulletin 

Vol. 125 contained 320 pages and the 43 papers covered a broad geographical spread with a strong accent 
on the Neotropics and Afrotropics. Although only two new taxa were described, a new species of 
Scytalopus tapaculo from Colombia and a new subspecies of Sylvia warbler from north-west Africa, the 
traditional Bulletin topics of taxonomy and nomenclature remained well represented, with a particularly 
comprehensive contribution on the nomenclature of Eduard Riippell's types from north-east Africa and 
several new generic designations being highlights. For papers published in 2005, the interval between 
receipt and publication was 6-18 months, averaging c. 12 months, which is a reasonable improvement on 
most recent years, although it is hoped to reduce 'waiting time' slightly further. The Hon. Editor is 
working on this issue as a medium-term objective. The Bulletin retains its popularity, with a total of 52 
new manuscripts being received in 2005. Of these, 16 were rejected, and the remainder are still being 
refereed. 

Grateftil thanks, as always, to referees who have given freely of their time and expertise, and to Eng- 
Li Green, of Alcedo Publishing, and Mike Dawson and team, of Crowes (our printers), for their support 
and efficiency. Thanks are also due to Mary MuUer who produced the index. Tony Statham produced the 
cover information and Club Announcements, and staff at the Natural History Museum, Tring, proffered 
much-valued support to the Hon. Editor. 

Bulletin — Institutional Subscriptions 

Subscriptions from Institutional Subscribers during 2005 totalled 116 (105 in 2004), with 27 in the UK 
and 89 overseas (28 different countries). Of the latter, 38 were from North America, 30 from Europe 
(including Russia), eight from Australasia, five from Africa, three from Asia and one from South 
America. 

Report of the Joint Publications Committee 

The BOC-BOU Joint Publications Committee (JPC) met twice in 2005. The Chairman of the committee 
is alternately appointed by the BOC and BOU. Nominated this year by the BOC, Tom Gladwin will chair 
the committee until April 2007. Bob Cheke has filled the vacancy, left by the late Janet Kear, as 
Commissioning Editor for all titles for which the committee is responsible. The Bird Atlas of Uganda was 



Club Announcements 87 Bull. B.O.C. 2006 126(2) 

published during the year and several other titles progressed towards publication. In response to 
requested changes, the proposed checklist The Birds of Singapore has been withdrawn by the author. 

Forthcoming Publications In the checklist series, The Birds of Sao Tome and Principe and Annobon 
will be published early in 2006. The Birds of Borneo and The Birds of Barbados continue to progress, 
and a manuscript is expected on The Birds of Crete. Priority will be given to publishing The Status of 
Birds in Britain and Ireland by the spring of 2008 as part of the BOU's 150th anniversary celebrations. 
The proceedings of the conference on Recent Avian Extinctions, which will be published in Bull. Brit. 
Orn. CI., will appear in 2006. 

Taxonomy The JPC has concerns about the liberal approach to taxonomic principles adopted by 
some proposed authors. It is seeking advice in order to make recommendations that authors should fol- 
low an established and authorised system, or otherwise with the JPC's prior consent. 

Acknowledgements The Committee is grateful to all who have contributed to its work and especial- 
ly to Bob Cheke as Commissioning Editor, and Steve Dudley for progressing and managing the titles in 
its charge. 

Membership 

Subscription rates were increased from 1 January 2005. As at 31 December 2005, there were 512 paid- 
up Members (539 in 2004).- 273 from the UK and 239 from overseas (44 countries). Overseas members 
comprise Europe (104), North America (60), Australasia (23), Africa (21), Asia (17) and South America 
(14). The Club welcomed 12 new Members, but mourned the death of Members E. D. H. Johnson 
(1970-2005), K. V. Thompson (1979-2004), R. S. R Fitter (1943-2005, Committee 1959-62, Vice- 
Chairman 1962-65, Chairman 1965-68), and A. J. Helbig (1994-2005). Ten resignations were received 
plus two who gave advance notice they would not renew for 2006, and 12 were removed under Rule 24, 
resuhing in a net decrease in membership of 27 during the year. 

Finance 

After a busy year for the Club, which saw the long-awaited publication of the Bird Atlas of Uganda, with 
its substantial printing costs, the Club's finances broke even in 2005. A loss of £2,584 on Unrestricted 
Funds was almost exactly matched by an offsetting surplus of £2,576 on Restricted Funds to give a net 
loss for the year of £8 before allowing for the changes in the market value of investments. 

Income for the year totalled £38,471, including £12,883 from subscriptions, which benefited from 
the increases that applied from 1 January 2005 and a consequent increase to £758 in the amount of tax 
recovered from Gift Aid and Deeds of Covenant. Investment income rose by £1,195 to £17,965. Of this 
sum, £10,410 came from the Herbert Stevens Fund, giving a yield on the year-end market value of 
4.05%. Interest on cash deposits amounted to £7,5 19, mostly from cash invested in charity deposit funds. 
Sales of BOC publications produced £1,100, with approximately half coming from sales of Why 
Museums Matter, but income from sales from BOC/BOU joint publications dropped by £2,200, as sales 
of previous year's Checklists declined and sales of the Uganda Atlas took off rather slowly in the second 
half of the year. 

Total expenditure, at £38,479, was well up on the previous year. Two main items accounted for most 
of this increase. Final production costs of the Uganda Atlas amounted to £11,331 and have been written 
off in full in the year, and there was an increase of £2,400 in the management fee paid to the BOU for 
the services of Steve Dudley who now handles sales of publications and assumed responsibility for all 
subscription matters part way through the year. Bulletin costs, including the honorarium paid to the 
Editor, showed only a slight increase to £13,932, but still run about £1,000 ahead of subscriptions. 

Investments 

Stock markets continued to improve during 2005, particularly in the second half of the year. As a result, 
the market value of the Club's investments in charity unit trusts held in the Herbert Stevens Fund showed 
an increase of £28,333 by the end of the year taking the value to £256,714, an increase of 12.4% over 12 
months and finally recovering from the substantial falls in 2001-02. The Barrington Trust Fund increased 
slightly to a year-end value of £1,162. As this now represents a very small part of the Club's finances, a 



Club Announcements 88 Bull. B.O.C. 2006 126(2) 

proposal will be made to the Annual General Meeting to close this fund and to transfer the money into 
the charity deposit account where it will earn approximately the same level of income. Total investments 
in the Balance Sheet are stated at £306,876, the additional £49,000 being the capital from the Clancey 
bequest, which continues to be held in a fixed-rate Building Society account. 

Reserves 

Of the Club's total resources, £20,000 remains in a Designated Unrestricted Fund towards the cost of 
future publications, two of which. The Birds of Borneo and The Status of Birds in Britain and Ireland are 
likely to be quite costly. The balance of the Uganda Atlas Fund was applied during the year against the 
final cost of publication and that fund is now closed. The two remaining funds, the Clancey bequest and 
the Publications Fund totalled £59,448 at the year-end and are held as backing for new developments for 
the Bulletin or additions to other Club publications, respectively. The investments that form a large part 
of the Unrestricted Funds built up from past legacies provide a regular source of income towards the 
Club's administrative expenses, whilst additional liquid funds are available to finance any future 
publications. This can involve substantial expenditure against a rather slow payback as expected with 
specialist publications that are highly valued, but fall within an inevitably limited market. 

Risks 

The Committee has reviewed the major risks to which the Club is exposed, particularly with regard to 
managing the Club's cash resources and overseeing the production of new publications within a 
controlled timetable. Investments are held in diversified portfolios within separately managed charity 
unit trusts providing an acceptable balance between security and risk. In particular, the Herbert Steven 
Fund is managed by three Trustees, who report regularly to the Committee. Additionally, the Joint 
Publications Committee considers proposals for further publications and puts forwards 
recommendations, bearing in mind the level of resources available. 

Trustees' Responsibilities 

Under the Charities Act 1993, the Trustees are required to prepare a statement of accounts for each 
financial year which gives a true and fair view of the state of affairs of the charity at the end of the 
financial year and of the incoming resources and application of resources in the year. In preparing the 
statement the Trustees are required to: 

Select suitable accounting policies and then apply them consistently; 

• Make judgements and estimates that are reasonable and prudent; 

State whether applicable accounting standards and statements of recommended practice have been 
followed, subject to any material departure disclosed and explained in the statement of accounts; 

• Prepare the financial accounts on the going concern basis unless it is inappropriate to presume that 
the charity will continue its operations. 

The Trustees are responsible for keeping proper accounting records which disclose with reasonable 
accuracy at any time the financial position of the charity and to enable them to ensure that any statement 
of accounts prepared by them complies with the regulations under secfion 41 (1) of the Charifies Act 
1993. They are also responsible for safeguarding the assets of the trust and hence for taking reasonable 
steps for the prevention and detection of fraud and any other irregularities. 

Approved and signed on behalf of the Trustees 

M. B. Casement 

Chairman Date: 25 April 2006 



Club Announcements 89 Bull. B.O.C. 2006 126(2) 

BRITISH ORNITHOLOGISTS' CLUB 
BALANCE SHEET— 31 December 2005 

Notes 2005 2004 

FIXED ASSETS 

Projection Equipment 
INVESTMENTS 

At market value 
CURRENT ASSETS 

Stock of publications 
Cash at bank and in hand 
Cash on deposit 
Prepayments 
Other debtors 



CURRENT LIABILITIES 

Subscriptions in advance 

Creditors falling due within one year 





£ 


£ 


£ 


£ 


2 




324 




486 


3 


100 

5,285 

108,980 

375 

1,707 

116,447 

(4,848) 
(2,878) 


306,876 

108,721 
415,921 


100 

4,404 

109,828 

363 

1,692 

116,387 

(4,142) 
(3.678) 


278,384 
108,567 




387,437 


4 




20,000 




20,000 


5 




336,473 
356,473 




310,565 
330,565 


6 




59,448 
415,921 




56,872 
387,437 



TOTAL ASSETS 

FUNDS 
Unrestricted 

Designated 
Other 

Restricted 



Approved and Signed on behalf of the Trustees 

MB Casement 

Chairman Date: 25 April 2006 

STATEMENT OF FINANCIAL ACTIVITIES— 3 1 December 2005 







2005 




2004 




Unrestricted 


Restricted 


Total 


Total 




£ 


£ 


£ 


£ 


VCOMING RESOURCES 










SUBSCRIPTIONS 










Members 


9,474 


— 


9,474 


6,970 


Institutional subscribers 


2,651 


— 


2,651 


2,300 


Income Tax recoverable 










under Gift Aid «& Deeds of Covenant 


758 

12,883 

97 


— 


758 

12,883 

97 


590 




— 


9,860 


DONATIONS 


— 


38 



Club Announcements 

INVESTMENT INCOME 

Herbert Stevens Trust Fund 
BaiTington Trust Fund 
Interest received 



SALES OF PUBLICATIONS 

Bulletin 

Other BOG publications 

Joint BOU/BOC publications 



90 



Bull. B.O.C. 2006 126(2) 



MEETINGS 
OTHER INCOME 

TOTAL INCOME 



10,410 


— 


10,410 


10,215 


36 


— 


36 


36 


4,722 


2,797 


7,519 


6,519 


15,168 


2,797 


17,965 


16,770 


156 


_ 


156 


160 


1,100 


— 


1,100 


831 


2,237 


— 


2,237 
3,493 
3,860 


4,475 


3,493 


— 


5,466 


3,860 


— 


4,127 


173 


— 


173 
38,471 


351 


35,674 


2,797 


36,612 



RESOURCES EXPENDED 

CHARITABLE EXPENDITURE 
MEETINGS 

Room and equipment hire, speakers' expenses, etc. 

Restaurant 
CONFERENCE SPONSORSHIP 
BOC BULLETIN 

Production, printing and distribution 
OTHER PUBLICATIONS 

Production costs 

Publicity, postage and packing 
ADMINISTRATION 7 

OTHER EXPENDITURE 

TOTAL EXPENDITURE 

SHORTFALL / EXCESS OF INCOME 
OVER EXPENDITURE 

Increase in value of investments 



TOTAL FUNDS brought 
forward at 1 January 2005 

TOTAL FUNDS at 3 1 December 2005 



1,588 
4,080 



13,932 



1,588 1,468 

4,080 4,119 

— 500 

13,932 13,821 



11,110 


221 


11,331 


7,762 


688 


— 


688 


945 


6,860 


— 


6,860 


3,807 


— 


— 


— 


180 


38,258 


221 


38,479 


26,602 


(2,584) 


2,576 


(8) 


10,010 


28,492 


— 


28,492 
28,484 


20,067 


25,908 


2,576 


30,077 


330,565 


56,872 


387,437 


357,360 


356,473 


59,448 


415,921 


387,437 



Club Announcements 91 Bull. B.O.C. 2006 126(2) 

NOTES TO THE ACCOUNTS— 31 December 2005 

1. ACCOUNTING POLICIES 

a) Basis of Accounts. The financial statements are prepared under the historical cost convention as 
modified by the inclusion of investments in the Herbert Stevens and Barrington Trust Funds at market 
values. They are also prepared in accordance with the Financial Reporting Standards for Smaller Entities 
and follow the recommendations in Accounting and Reporting by Charities: Statement of Recommended 
Practice (revised 2005). 

b) Investments and Cash Deposits. The Herbert Stevens and Barrington Trust Funds are invested in 
quoted charity unit trusts and included as investments in the Balance Sheet at year-end market values. 
Income from these funds and from cash deposits shown in the Balance Sheet under Current Assets is 
included in Incoming Resources in the Statement of Financial Activities on a receipts basis. The Clancey 
Bequest is held in a fixed term, Building Society deposit account which forms part of the total of 
investments in the Balance Sheet. Interest on this deposit account is brought into the Statement of 
Financial Activities on an accruals basis. 

c) Subscriptions. Subscriptions for the current year and any arrears are included in Incoming Resources 
in the Statement of Financial Activities. Subscriptions received in advance are carried forward in the 
Balance Sheet as Current Liabilities. 

d) Expenditure is accounted for on an accruals basis. 

e) Depreciation. Depreciation of fixed assets is calculated to write off their value over their expected 
useful lives at an annual rate of 25% on cost. 

f) Publications. The cost of publications is written off in the Statement of Financial Activities as 
incurred except for a nominal stock value of £100 carried in the Balance Sheet. 

2. FIXED ASSETS 2005 

Projection Equipment 
Cost at 1 January and 3 1 December 2005 
Accumulated depreciation at 1 st January 2005 
Charge for the year 
At 3 1 December 2005 
Net Book Value: At 3 1 December 2005 
At 31 Decemebr 2004 

3. INVESTMENTS— at market value 

UNRESTRICTED FUNDS 

Herbert Stevens Trust Fund 
Barrington Trust Fund 

RESTRICTED FUNDS 

Clancey Bequest 49,000 49,000 



£ 




648 




162 




162 




324 




324 




486 




2005 


2004 


£ 


£ 


256,714 


228,381 


1,162 


1,003 



306,876 278,384 



All investments are held in the UK. 



4. UNRESTRICTED DESIGNATED FUND 2005 

for future publications £ 

Balance at 1 January 2005 20,000 

Designated during the year -- 

Balance at 3 1 December 2005 20,000 



Club Announcements 



92 



Bull.B.O.C. 2006 126(2) 



HERBERT 


BARRINGTON 




STEVENS 


TRUST 




TRUST FUND 


FUND 


TOTAL 


£ 


£ 


£ 


228,381 


1,003 


310,565 


28,333 


159 


28,492 


( 2,584) 







5. OTHER UNRESTRICTED FUNDS 

GENERAL 
FUND 

£ 
Balances at 1 January 2005 81,181 

Increase in value of investments during year — 
Shortfall of income over expenditure ( 2,584) 

Balances at 3 1 December 2005 

6. RESTRICTED FUNDS 



Balances at 1 January 2005 
Expenditure during the year 
Interest received 

Balances at 3 1 December 2005 



a . The Clancey bequest was donated by the late Dr P. A. Clancey with the request that it should be used 

to support and enhance the Club's Bulletin, 
b . The Publications Fund is available to finance Club publications other than regular issues of the 

Bulletin, 
c . The Bird Atlas of Uganda Fund was earmarked to assist with the production and publication of the 

Atlas and has now been fully applied for that purpose. 



78,597 256,714 


1,162 


336,473 


PUBLICATIONS 
CLANCEY FUND 
BEQUEST 

£ £ 

52,528 4,132 
(221) (221) 
2,601 187 


BIRD ATLAS 

OF UGANDA 

FUND 

£ 

212 

9 


TOTAL 

£ 

56,872 

2,797 


55,129 4,319 


— 


59,448 



7 . ADMINISTRATION EXPENSES 



Club's share of rental of storage unit 

Audit and Independent Examination fees 

Depreciation 

BOU administration services 

Other administration expenses 



2005 


2004 


£ 


£ 


916 


916 


550 


550 


162 


162 


3,080 


642 


2,152 


1,537 


6,860 


3,807 



8. REIMBURSEMENT OF EXPENSES 

Trustees do not receive any remuneration or any reimbursement of costs they may incur in attending 
Trustee meetings. They are reimbursed for any other expenses incurred on behalf of the Club. The total 
amount reimbursed during the year was £1,072 (2004 £1,576). 



Club Announcements 93 Bull. B.O.C. 2006 126(2) 

INDEPENDENT EXAMINERS REPORT TO THE TRUSTEES OF 
THE BRITISH ORNITHOLOGISTS' CLUB 

I report on the accounts of the Club for the year ended 31 December 2005, which are set out on pages 
89-92. 

Respective responsibilities of trustees and examiner 

The charity's trustees are responsible for the preparation of the accounts. The charity's trustees consider 
that an audit is not required for this year (under section 43(2) of the Charities Act 1993 (the 1993 Act)) 
and that an independent examination is needed. 
It is my responsibility to: 

Examine the accounts (under section 43(3)(a) of the 1993 Act); 
• To follow the procedures laid down in the General Directions given by the Charity Commissioners 

(under section 43(7)(b) of the 1993 Act); and 

To state whether particular matters have come to my attention. 

Basis of independent examiners' report 

My examination was carried out in accordance with the General Directions given by the Charity 
Commissioners. An examination includes a review of the accounting records kept by the charity and a 
comparison of the accounts presented with those records. It also includes consideration of any unusual 
items or disclosures in the accounts, and the seeking of explanations from you as trustees concerning any 
such matters. The procedures undertaken do not provide all the evidence that would be required in an 
audit and, consequently, 1 do not express an audit opinion on the view given by the accounts. 

Independent examiners' statement 

In connection with my examination, no matter has come to my attention: 

(1) which gives me reasonable cause to believe that, in any material respect, the requirements: 

• to keep accounting records in accordance with s41 of the 1993 Act; and 

• to prepare accounts which accord with the accounting records and to comply with the 
accounting requirements of the 1993 Act have not been met; or 

(2) to which, in my opinion, attention should be drawn in order to enable a proper understanding of the 
accounts to be reached. 

Alan Peal 

Alan Peal ACA — Principal 
Porritt Rainey 
Chartered Accountants 

25 April 2006 



Thomas M. Donegan & Blanca Huertas 94 Bull. B.O.C. 2006 126(2) 

A new brush-finch in the Atlapetes latinuchus 

complex fii'oni the Yariguies Mountains and 

adjacent Eastern Andes of Colombia 

by Thomas M. Donegan & Blanca Huertas 

Received 28 May 2005; final revision received 13 February 2006 

The brush-finches Atlapetes are Neotropical passerines which achieve greatest 
diversity in the Andes and whose taxonomy and ecology have received much recent 
attention. Atlapetes shows a high degree of geographical variation, with various 
forms restricted to particular elevations, mountains or slopes, but few cases of true 
sympatry. The group is therefore ideally suited to studies of the patterns and 
mechanisms of speciation (Remsen & Graves 1995a, Garcia-Moreno & Fjeldsa 
1999). Even recently, localised taxa have been described, both at species and 
subspecies level (e.g. Fitzpatrick 1980, Remsen 1993, Valqui & Fjeldsa 1999), and 
a species thought possibly extinct was rediscovered (Agreda et al. 1999). 

Serrania de los Yariguies and the Eastern Cordillera 

The Eastern Cordillera (Eastern Andes) is one of Colombia's three principal 
mountain ranges, extending from dpto. Cauca (01°N), to the Serrania de Perija, on 
the Caribbean coast (11°N). It is characterised by a varied habitats, with slopes 
bordering Amazonia, the llanos and Magdalena Valley, humid and dry regions, 
plateaux, steep slopes and wetlands. The Cordillera's cool climate and high- 
elevation savannas have long been subject to human development. It is also one of 
the world's major centres of avian endemism, the Colombian East Andes Endemic 
Bird Area (Stattersfield et al. 1998). 

The Serrania de los Yariguies (dpto. Santander) is an isolated western spur of the 
East Andes, rising to c.3,400 m and isolated from the rest of the cordillera to the 
north and east by the Sogamoso Valley, and to a lesser extent to the south by 
depressions associated with the rios Horta, Quirola and Opon and their tributaries. 
A collection of 60 bird species was made below 1,000 m near San Vicente de 
Chucuri in November 1956 (Borrero & Hernandez 1957), and a few specimens were 
taken elsewhere on the massif, generally on the drier eastern slope (e.g. Romero 
1983), but the humid western slope and highest elevations were very poorly known 
ornithological ly prior to our field work (for general results see Donegan & Briceiio 
2005, Donegan & Huertas 2005, Donegan & Avendano 2006, Huertas & Donegan 
2006). Amongst the birds we recorded was one which initially appeared to be of the 
'Northern Rufous-naped Brush-fmch' or 'Yellow-breasted Brush-fmch' complex, 
A. latinuchus. 

Paynter (1978) specifically commented on the absence of records of A. 
latinuchus (then classified as A. rufinucha) from the central part of the Eastern 
Cordillera, noting "While it is present in the Sierra de Perija, a northward extension 



Thomas M. Donegan & Blanca Huertas 95 Bull. B.O.C. 2006 126(2) 

of the Eastern Andes on the Venezuela-Colombia border, it has not been found in 
the Eastern Andes except for a few dubious records from Bogota {A. r. simplex, 
known only from native Bogota specimens)'. The range of .4. /. simplex was 
subsequently thought to comprise the central eastern plateaux and perhaps the east 
slope of the Eastern Andes between Lago de Fuquene (dpto. Cundinamarca) and 
Soata (dpto. Boyaca) (Borrero & Olivares 1955, Hilty & Brown 1986, Ridgely & 
Tudor 1989). However, such records and reported 'variations' in the plumage of ^. 
/. simplex derive from immatures and subadults in collections, a failure to compare 
types and new material, and the existence of two taxa from different regions. 

Methods 

The Colombian EBA (Evaluation of Biodiversity in the Andes) Project is an 
ongoing research and conservation initiative, supported by Fundacion ProAves, that 
has conducted Rapid Biodiversity Assessments of remote and/or unstudied sites 
since the mid 1990s. 

We studied ten primary-forest sites on both slopes of the Serrania de los 
Yariguies at elevations of 100-3,200 m between January 2003 and January 2006. 
Each site was subject to 4-6 days' field work using mist-netting (up to 220 m of 
mist-nets) and non-systematic observations including sound-recording and 
playback. On 4-8 January 2004, we visited a remote site, at 2,400-2,450 m, on the 
west slope, accessed from Cantagallos, in San Vicente de Chucuri municipality, to 
Finca Santo Domingo (2,250 m), from where we reopened a disused trail to the 
ridge at 2,450 m. 

On 6 January 2004, together with Elkin Briceflo, we trapped an Atlapetes which 
showed features of the A. latinuchus complex, e.g. a rufous cap, black mask, yellow 
underparts and an indistinct moustachial (Fig. lA-B), but differed from all other 
populations in its jet black mantle, tail and wings, and from some taxa in the group 
by the lack of a white speculum. A. latinuchus is unknown from the north-central 
Eastern Cordillera (Hilty & Brown 1986), with the distinctive form A. I. nigrifrons 
to the north and paler-backed A. I. simplex postulated to occur further south in the 
Cordillera. Given this, we considered it probable that the bird represented an 
undescribed taxon. Due to poor weather conditions for photography, we retained the 
bird overnight for further study but, unusually for an Atlapetes, it died overnight. 
The bird was prepared in the field as a flat skin. 

Comparison of our photographs and wing- and tail-feathers from the bird with a 
'Bogota' A. I. simplex in the Natural History Museum (NHM; Tring) revealed 
various differences between them, with A. I. simplex having a grey/brown back, 
paler red crown, small white speculum and larger yellow supraloral spot (Fig. IC). 
Further research revealed additional specimens of the black-backed form from the 
Virolin area, dpto. Santander, on the west slope of the main Eastern Cordillera, c.60 
km south-east of Serrania de los Yariguies. On 8 January 2005, at Lepipuerto, on the 
upper rio Chimera, El Carmen/Simacota municipality (06°28'N, 73°28'W; 
2,900 m), TD and Martin Donegan observed and sound-recorded a pair foraging in 



Thomas M. Donegan & Blanca Huertas 96 



Bull. B.O.C. 2006 126(2) 




Figures 1 A-B. Holotype of ^. /. yariguierum (Blanca Huertas/EBA Project) 




Figure IC. A. I. simplex (=A. I. spodionotus) neotype (Thomas Donegan, © Natural History Museum, 
Tring). Note difference in contrast between upperparts and mask colour compared to the holotype, the 
very small but visible white speculum and relatively strong malar, each consistent with^. /. spodionotus. 



Thomas M. Donegan & Blanca Huertas 97 Bull. B.O.C. 2006 126(2) 

paramo. Further observations and the capture of another bird (from which a blood 
sample was obtained) were made in July 2005, during 'Proyecto YARE', by 
ourselves and Jorge Avendaiio in paramo at Filo Pamplona, Galan municipality 
(06°38'N, 73°24'W; 3,200 m), on the east slope of the massif 

Type material ofA.L sitnplex 

Project Biomap provided data from all museums holding Colombian specimens. We 
also searched an online database (www.biologie.uni-ulm.de) of all major German 
museums, but were unable to locate the two A. I. simplex specimens mentioned in 
the type description (Berlepsch 1888). All remaining specimens from the Berlepsch 
collection are now held in Frankfurt, except a few specimens in Warsaw 
(F. Steinheimer in lift. 2005), neither of which institutions possess any A. I. simplex 




Figure ID. Close-up of mantles and crown of (left to right) adult A. I. simplex neotype ('Bogota', 
Colombia = ^. /. spodionotus), subadult^. /. elaeopronis (Central Andes, Antioquia, Colombia), adult ^4. 
/. comptus (southern Ecuador and northern Peru), and adult A. I. spodionotus (Ecuador and southern 
Colombia). (Thomas Donegan, © Natural History Museum, Tring). Note contrast between the grey 
upperparts and black face on A. I. simplex compared with much darker A. I. elaeoprorus, and the paler 
crown of A. I. simplex compared to A. I. elaeoprorus. A. I. yariguierum averages blacker than A. I. 
elaeoprorus with essentially no contrast between the mask and upperparts in adults. Note also similarity 
between 'A. I. simplex' and A. I. spodionotus. NB: adult male A. I. elaeoprorus has essentially black 
upperparts (as in tail of the subadult), not black admixed olive. 



Thomas M. Donegan & Blanca Huertas 98 Bull. B.O.C. 2006 126(2) 

(G Mayr in litt. 2005, N. Krabbe in lift. 2006). It seems probable that the A. I. 
simplex types were either destroyed or lost during World War II (G. Mayr in litt. 
2005). The A. I. simplex at NHM is a male, apparently adult, originally from the 
Berlepsch collection. It too is dated 1888, and the label is annotated 'agrees with 
types' in Berlepsch 's hand. The plumage, particularly the trace of a small white 
speculum, is consistent with Berlepsch's (1888) brief description (although some 
apparent adult/immature variation is noted, with the adult somewhat darker). This 
specimen, like the types, is labelled 'Bogota'. Thus, we assume this specimen to be 
typical of the form described by Berlepsch as A. I. simplex, and propose to treat it 
as a neotype of ^4. /. simplex. 

A. L simplex is a junior synonym oiA, L spodionotus 

The A. I. simplex neotype is indistinguishable from A. I. spodionotus of the main 
Andean range in southern Colombia and northern Ecuador. 'Bogota' trade skins 
originated from as far away as Panama or Ecuador and several taxa (including the 
recently described Bogota Sunangel Heliangelus zusii) are known only from such 
material (see Graves 1993). A. I. simplex was described with, and appeared in the 
same consignment as, the antbird taxon Myrmeciza longipes boucardi, a form now 
known to be restricted the upper Magdalena Valley of Colombia, from dpto. Tolima 
to the headwaters at 1,700 m (Hilty & Brown 1986). It seems feasible that A. I. 
simplex skins may originate from slightly further south, where A. I. spodionotus is 
present. Other taxa until recently known in Colombia only or principally from 
Bogota skins, e.g. Band-bellied Owl Pulsatrix melanota. Black-streaked Puffbird 
Malacoptila fulvogularis and Red-billed Parrot Pionus sordidus, have recently been 
found there (Salaman et al. 2002). The case for A. I. simplex being distinct from A. 
I. spodionotus was not made by Berlepsch (1888), who focused on the differences 
from A. I. elaeoprorus of Antioquia. It seems that A. I. simplex is better treated as a 
junior synonym of ^. /. spodionotus (Sclater & Salvin 1879). 

Description 

With the status of ^. /. simplex resolved, questions concerning the dark-backed 
populations of ^. latinuchus in the Serrania de los Yariguies and adjacent slope of 
the Eastern Cordillera is clarified, and some supposed 'variations' in plumage of 
birds from this region can be discarded. This Eastern Cordillera population differs 
from all other described Atlapetes taxa. We propose to name it: 

Atlapetes latinuchus yariguierum subsp. nov. 

Yariguies Brush-finch 

Gorrion-montes de los Yariguies 

Holotype Adult male, no. ICN-34016 of the ornithological collection of the Instituto 
de Ciencias Naturales, Universidad Nacional de Colombia (ICN), Bogota, 
Colombia (Fig. la). Prepared by TD, on 7 January 2004, at Alto Cantagallos, 



Thomas M. Donegan & Blanca Huertas 99 Bull. B.O.C. 2006 126(2) 

Serrania de los Yariguies, San Vicente de Chucuri municipality, dpto. Santander, 
Colombia (06M9'N, 73°22'W). Study specimen produced from flat skin by F. G 
Stiles. The locality is at 2,400 m on the west slope of the Yariguies massif, Eastern 
Andes, in lower montane cloud forest. Tissue samples and stomach contents are held 
at the Instituto Alexander von Humboldt's molecular laboratory, Cali, Colombia. 

Paratypes and other material We examined specimens of all A. latinuchus taxa 
(except^. /. chugurensis) in the following institutions: ICN (Bogota), NHM (Tring), 
University Museum of Zoology, Cambridge, UK (UMZC), Instituto Alexander von 
Humboldt, Villa de Leyva, Colombia (lAVH), Museo de la Universidad de la Salle, 
Bogota, Colombia (MLS), Museo de Historia Natural, Universidad Industrial de 
Santander, Bucaramanga, Colombia (UIS) and Museum National d'Histoire 
Naturelle, Paris, France (MNHN). Digital photographs of all A. latinuchus 
specimens in the Coleccion Omitologica Phelps, Caracas, Venezuela (COP) and 
some in the Field Museum of Natural History, Chicago, USA (FMNH) were also 
inspected (see Appendix 1). We assign the following paratypes: 

Adult sex unknown, no. UIS 1412, collected by V H. Serrano, on 23 August 
2001, at Santa Helena (06"00'N, 73^09'W), between Charala and Duitama, Boyaca, 
at 2,400 m, on the west slope of the Eastern Andes. Subadult female, no. ICN- 
25111, prepared 28 November 1978 (collector unknown), from Charala, dpto. 
Santander (c.06°05'N, 73°12'W) on the west slope of the Eastern Andes. Subaduh 
(probably female), DNA sample and photographs taken on 1 2 July 2005 by TMD & 
J. Avendaiio, at Filo Pamplona, Serrania de los Yariguies (06°38'N, 73°24'W; 
3,200 m); DNA sample deposited at UIS. 

The following juveniles also appear to be of the new taxon but are not assigned 
as paratypes: juvenile male, no. ICN-3199, collected by A. Olivares, on 7 January 
1953, at Soata, Alto de Onzaga, Boyaca, (c.06°35'N, 72°07'W) on the east slope of 
the East Andes (juvenile plumages are poorly known, making this is a tentative 
identification); juvenile of unknown sex, no. ICN- 10322, collected by A. Olivares 
& P. Bemal, on 9 July 1961, in Vereda Ruparita, 2 km east of Arcabuco, Boyaca, 
(c.05"73'N, 73°45'W) on the west slope of the East Andes. 

The following specimens, of which we have seen digital photographs, are also 
apparently of the new taxon but are not assigned as paratypes: subadult male, no. 
FMNH-220606, collected by Kjell von Sneidem, in 1950, above Chiquinqira, 
Boyaca (c.05°37'N, 73°50'W) on the east slope of the East Andes; juvenile, no. 
FMNH-220607, taken by the same collectors, from the same locality, on the same 
dateasICN-10322. 

Diagnosis A typical Atlapetes with various characteristics of the A. latinuchus 
complex, including dark upperparts and yellow underparts, a rufous crown, 
(concealed) white speculum, black mask and indistinct moustachial (Hilty & Brown 
1986), but differing from others within the A. rufinucha complex by its sharply 
demarcated rufous cap (at least in adults), yellow malar, slight melanism on the 



Thomas M. Donegan & Blanca Huertas 



100 



Bull. B.O.C. 2006 126(2) 



flanks, and white speculum (Garcia-Moreno & Fjeldsa 1999). Analysis of molecular 
data shows it to be related to ^. /. elaeoprorus and A. I. spodionotus (see below). 

A. I. yariguierum differs from all forms of A. latinuchus in its uniform jet black 
mantle, tail, and wings (with no olivaceous or greyish tones, except in juveniles, or 
white speculum), and virtually no contrast between the black mask and upperparts 
(though some adult A. I. elaeoprorus approach this). A 'rather greyish back' was 
considered diagnostic of the A. latinuchus group by Garcia-Moreno & Fjeldsa 
(1999). Adult A. I. yariguierum is notably darker rufous on the crown than other A. 
latinuchus taxa (some adult A. I. elaeoprorus approach it), and is further 
distinguished by the lack of a visible speculum, obvious supraloral spot, dark 
forehead markings, strong malar or white/cream on the nape. Similarities between 
A. I. elaeoprorus and A. I. yariguierum perhaps indicate previous contact between 
taxa of the northern Central and Eastern Cordilleras, also evidenced amongst 
subspecies of Grallaricula nana, Phaethornis syrmatophorus and Basileuterus 




Figure 2. From left to right (youngest to oldest): juvenile apparent^. /. yariguierum (Soata, Boyaca, east 
slope of East Andes); juvenile A. I. yariguierum (Arcabuco, Boyaca, west slope of East Andes); subadult 
paratype A. I. yariguierum (Virolin, Santander, Colombia, west slope of East Andes), and adult holotype 
A. I. yariguierum (Serrania de los Yariguies, Santander, Colombia) at ICN-UN (Thomas Donegan). Note 
black tails of all birds. Ages ascribed on basis of plumage as no skull ossification data on specimen labels 
(except holotype). 



Thomas M. Donegan & Blanca Huertas 101 Bull. B.O.C. 2006 126(2) 

tristriatus (TMD unpubl). However, A. I. elaeoprorus always has a large white 
speculum and large yellow supraloral (unlike the almost indiscernible short line of 
yellow feathers below the rufous cap in A. I. yariguierum). Juvenile A. I. 
elaeoprorus has the back tinged dark greenish, whereas in ^. /. yariguierum it is 
tinged dark brownish. 

A. I. yariguierum differs from A. I. spodionotus in its combination of black 
mantle and tail, deeper rufous crown and reduced yellow feathering at the bill base. 
A report of the supraloral being variable in A. I. simplex (Paynter 1978) appears 
based on differences between the types and recent Eastern Cordillera material. Some 
A. I. spodionotus (including the A. I. simplex in Tring) show traces of a visible 
speculum in the folded wing. In all A. I. yariguierum observed in the field or 
museum, the speculum is invisible on the folded wing, though this feature could be 
visible, especially when moulting. In A. I. spodionotus, where the speculum extends 
almost to the greater coverts, it is visible in some individuals and not in others 
(Paynter 1978). 

A. I. yariguierum is separable from the A. schistaceus group by its yellow 
underparts and lack of strong moustachial or supraloral; from A. albofrenatus taxa 
by the absence of a well-defined moustachial and its black (not olive-green) mantle; 
from A. I. nigrifrons and A. melanocephalus by the lack of a black upper throat or 
forehead, pale moustachial or grey lores, and from the latter by its rufous cap; and 
from A. pallidinucha taxa in its uniform dark rufous crown and nape in adults. 
Morphometries of these taxa are presented in Appendix 3. 

Description of holotype Capitalised colour nomenclature and numbers from 
Munsell Color (1977) and Munsell Color (2000). Aduh male with skull 100% 
ossified (mass 22.6 g). Testes 8 mm x 5 mm (apparently somewhat enlarged with 
small cloacal protuberance in life) and little subcutaneous fat. Stomach contents 
included various small pieces of Coleoptera exoskeleton. Twelve rectrices, nine 
primaries, six secondaries and three tertials, typical of the Emberizinae. Flight- 
feathers fresh, with rectrices full (except fifth rectrix from left, 95% emerged), 
suggesting recent completion of mouh. Emarginated pp6-9 (slight emargination on 
p5), with rounded wing point pp4-6 and primary notches absent (following Proctor 
& Lynch 1993). Max. flattened wing (following Svensson 1992) 77 mm, tail 80 mm, 
tarsus 25 mm, culmen to skull 16 mm. Crown dark rufous (closest to P, lOR: 4/8); 
facial mask jet black; mantle, wing-coverts, alula, flight-feathers, rump and rectrices 
closest to black (Gley 1: 2.5/2.5 but darker; cf. greyish brown, closest to lOYR 3/1 
or 2/2 of 'A. I. simplex' and A. I. spodionotus), with virtually no contrast between 
mask and back. Very narrow line of six short Yellow (5Y 8/12) feathers below 
rufous crown, at base of bill. Throat, malar, breast and belly uniform Yellow (5Y 
8/12), becoming slightly yellower on belly and washed slightly darker on flanks. 
Indistinct darkish moustachial extends c.l6 mm from bill. Underside of carpal 
Yellow (5Y 8/12) with small blackish spots. Base of pp4-6 concealed under greater 
coverts frontally White (Gley 1 8/1). Each except outermost primary and innermost 



Thomas M. Donegan & Blanca Huertas 



102 



Bull. B.O.C. 2006 126(2) 



five secondaries distally near White (Gley 1 8/1). Bare parts: mandible uniform 
black; legs horn, with feet soles yellowish grey; iris dark rufous. 

Variation in the series Plumage and biometrics of the adult paratype essentially 
very similar to those of the holotype. The subadult paratype at UIS has a slightly 
paler rufous cap (c.2.5YR 4/8, but still darker than adult A. I. simplex! spodionotus). 



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Figure 3. Map showing sites at which A. I. yariguierum has been reported (blue squares; unfilled squares 
= unconfirmed records). All A. I. yariguierum records post-date Paynter (1972). Also shown are known 
ranges of /i. /. elaeoprorus (unfilled red circles); A. I. caucae (purple-filled circles); and^i. /. spodionotus 
(unfilled pink ellipses) in Colombia, based on specimens, published literature, data fi-om Project Biomap 
and additional sight records reported to the authors. Apparent gaps in distribution are partly due to 
observational lacunae, sometimes exacerbated by deforestation, and partly due to ranges of congeneric 
ecological competitors (see Remsen & Graves 1995). The southernmost record of ^. /. elaeoprorus in the 
West Andes and the northernmost, from the Serrania de San Lucas, are from data supplied by Project 
Biomap. Both require confirmation. Observadons from the East Andes in Cauca are of birds with a strong 
white speculum (J. Idrobo in litt. 2005) and therefore appear more likely to be of A. I. caucae than A. I. 
spodionotus. Of this group, A. I. yariguierum emerges as the taxon known from the largest number of 
localities in Colombia. 



Thomas M. Donegan & Blanca Huertas 



103 



Bull. B.O.C. 2006 126(2) 



kHz 
5 

4 

3- 
2- 
1 - 



- \ AtlapelAtlapetes 



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iiiliirtiiiliiiiiiiriii ' i lyiiJii 



1 s 




kHz 
5H 
4 
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kHz 




S8.975 



Figure 4. Vocalisations of ^. /. yahguierum recorded at Lepipuerto, Serrania de los Yariguies (top left and 
middle). Sonograms oi A. I. yahguierum produced by Richard Ranft (British Library) using Avisoft Pro 
software fi-om recordings by TMD. In each case, A. I. yahguierum vocalisations are presented together 
with the most similar recordings of ^. /. spodionotus from Yanacocha, Pichincha, Ecuador (00°08'S, 
78°35'W). Sonograms for A. I. spodionotus were produced by TMD using Raven Lite software, from 
recordings by J. V. Moore in Krabbe et al. (2001). 



Thomas M. Donegan & Blanca Huertas 104 Bull. B.O.C. 2006 126(2) 

both black and brownish (particularly at the base of the rump) mantle feathers, but 
the tail- and flight-feathers and most of the upperparts similar to adults. The 
demarcation between the rufous crown and dark back is less sharp in non-adults of 
A. I. yariguierum, a feature of other ^. latinuchus taxa (birds in Vellinga et al. 2004 
are probably an example of this). ICN-3199 is similar to the subadult paratype, but 
has more brownish and fewer black feathers, a paler crown and less obvious 
demarcation between the crown and upperparts. It has a very pale rufous nape 
(approaching A. pallidinuchd) and more brown and fewer black feathers in the 
upperparts. The juvenile at FMNH has a darker crown and nape but similar 
upperparts. All specimens in the series have a jet black tail (as does A. I. elaeoprorus 
at all ages). The ICN series appears in Fig. 2. We cannot eliminate the possibility 
that A. I. yariguierum may involve two taxa or clinal variation: it seems feasible that 
those from Yariguies are on average slightly darker overall with less yellow on the 
supraloral than birds from the main cordillera. The pale-naped bird from the east 
slope in Soata also requires further investigation. 

Vocalisations We sound-recorded a pair observed at Lepipuerto (Fig. 4). The call is 
typical of the genus: a short series of high-pitched, rising and falling whistles at 
c.3-5 kHz: wiu-wiu, wiu-weeu-weeu, wee-weeu-weeu and wee-wiu-weeu-weeeu. 
TD also once heard a 'stuttering' finch song, probably^. /. yariguierum and similar 
to A. latinuchus in Ecuador, but this was not sound-recorded. Recordings are 
deposited at the British Library (no. 142861), London. 

Published sound-recordings of ^4. /. spodionotus from Ecuador suggest that ^. 
latinuchus taxa, like other oscines, possess a wide repertoire. Small possible 
differences between A. I. spodionotus and A. I. yariguierum are: A. I. yariguierum 
calls appear simpler and include notes that increase and decrease in pitch more 
uniformly (straight line on the sonogram), and which decrease in pitch more slowly, 
than in A. I. spodionotus. With just two recordings of A. I. yariguierum, it is 
impossible to know whether calls are significantly different from A. I. spodionotus, 
which has very varied calls (Krabbe et al. 2001). Vocalisations are poorly known for 
Colombian Atlapetes, begging additional research which may have taxonomic 
implications. 

Distribution It is probable that A. I. yariguierum is restricted to high elevations in 
the Serrania de los Yariguies and adjacent Eastern Andes (Fig. 3). We are aware of 
the following records: Galilea, Reserva Biologica Chachalu (06°05'N, 73^09'W; 
2,350 m, the paratype locality), trapped 30 October 2003 (weight 30 g) and 27 
November de 2003 (weight 28.7 g), with three sight records (D. A. Rincon G). 
Rogitama, dpto. Boyaca (c.05M7'N, 73°3rW), south-west of Chachalu, where 
frequently observed and an adult photographed (R. Chavarra C). Soata, Boyaca (see 
Paratypes), in oak forest at c.2,900-3,100 m (O. Cortes), El Talisman, Serrania de 
los Yariguies (c.06"5 1 'N, 73°22' W) photographed in February 2006 at 2,050 m (M. 
Sharp & C. Turner). 



Thomas M. Donegan & Blanca Huertas 105 Bull. B.O.C. 2006 126(2) 

The following records also probably involve A. I. yariguierum but require 
confirmation: Finca El Diviso, dpto. Santander (07°08'N, 73°02'W; 1,850m), on 30 
December 2004 (D. A. Rincon G); Mesa de los Otero, Curiti municipality, dpto. 
Santander (06°37'N, 73°00'W, 2,100-2,250 m), on 19 November 2004 (J. Parra B.); 
Santuario de Fauna y Flora Iguaque, dpto. Boyaca (05°40'N, 73°27'W, 
3,045-3,065 m) (C. D. Cadena & J. E. Zuluaga); and Sutamarchan, Serrania de 
Merchan, dpto. Boyaca (c.05°37'N, 73°38'W), in April 2003 (M. A. Echeverry). 

Overall, a pattern of fairly continuous distribution within humid forests of the 
central Eastern Andes, principally on the west slope but also apparently on the east 
side, emerges. To date, A. I. yariguierum has been recorded at 1,800-3,200 m, being 
apparently at least partially replaced lower by Yellow-throated Brush-fmch A. 
gutturalis (recorded at 2,000 m on the west slope of Serrania de los Yariguies) and 
Moustached Brush-fmch A. albofrenatus (recorded at 2,000 m on both slopes of 
Serrania de los Yariguies). In the main cordillera, it is at least partially replaced at 
high elevations by Slaty A. schistaceus and Pale-naped Brush-finches A. 
pallidinucha, though neither of these has been recorded in the Yariguies range in 
surveys almost to the peak, where A. I. yariguierum is present. Elevational and 
latitudinal distributions of these taxa are described and discussed in Remsen & 
Graves (1995a). The range of ^. /. yariguierum is shared by other endemics, e.g. 
Gorgeted Wood-quail Odontophorus strophium and Mountain Grackle 
Macroagelaius subalaris, which are both Critically Endangered (BirdLife 
International 2004). 

Etymology Our name honours the Yariguies indigenous people and the massif that 
bears their name, expressed as a fifth declension feminine Latin noun yariguies, and 
declined in the genitive plural. Serrania de los Yariguies is often labelled on maps 
as Serrania de los Cobardes (Mountains of the cowards), which derives from the 
colonial era. The Yariguies people resisted the Spanish, although was doomed to fail 
in the face of a technologically superior military power. Rather than submit, the 
Yariguies committed mass suicide. The Spanish viewed this not as an act of dignity, 
but of cowardice. Colombian governmental agencies and NGOs are endeavouring 
to remove the insult from official materials, and Serrania de los Yariguies is now the 
preferred name. The region is rich in archaeological sites and indigenous art, which 
to date have gone largely unrecognised due to the historic security situation and 
difficulty of access. 

Ecology 

The holotype was captured in a treefall gap with successional vegetation, c.l km 
from human-modified habitat, in forest on a west-facing slope overlooking the 
Magdalena Valley, characterised by a canopy of 8-14 m, dense understorey and 
epiphytic growth, persistent fog or rain, little insolation (less than 30 minutes daily 
during field work), and 30-50° slope. Observations at Lepipuerto were made in 
pristine primary treeline and paramo habitat c.30 km from any human population. 



Thomas M. Donegan & Blanca Huertas 106 Bull B.O.C. 2006 126(2) 

and also subject to very high levels of precipitation, with torrential rain lasting 
several hours each day during the 'dry' season, and frequent ground-level cloud 
cover. The pair observed here foraged in small tree-like shrubs on an exposed slope. 
Observations at the third site in Serrania de los Yariguies were also in pristine 
treeline and paramo habitat far from human populations. All three sites are within 
the same tract of forest, this being one of the largest such forests in northern 
Colombia. Successional primary forest, at the type locality, and fairly open 
vegetation of the Yariguies paramo appear to be the natural habitat of ^. /. 
yariguierum. A. latinuchus taxa regularly invade secondary habitats (Remsen 1993, 
Hilty, 2003). The new taxon also appears to do so, and indeed seems commoner in 
such habitats than in primary forest in the main Eastern Andes. 

Moult 

The very fresh plumage and emergent rectrix of the holotype indicate it had recently 
completed a moult, whilst the enlarged testes and cloacal protuberance indicate pre- 
breeding moult, and that this form's breeding season coincides with the wettest part 
of the year, in February- April (the main breeding season in the adjacent Central 
Andes of the Magdalena Medio: Cuervo et al. 2001). 

Discussion 

Garcia-Moreno & Fjeldsa (1999) recently re-evaluated species limits within 
Atlapetes using molecular data. The taxonomy of the group was previously based on 
morphological analyses (Paynter 1972, 1978), with the taxa A. schistaceus and A. 
rufinucha delimited largely on the presence of grey {schistaceus group) or yellow 
underparts {rufinucha; pileatus group). Thereafter, Remsen & Graves (1995a) 
suggested that several geographically close taxa with alternating grey and yellow 
underparts may be more closely related to one another than to more geographically 
distant taxa with similar underparts coloration. Garcia-Moreno & Fjeldsa's 
phylogeny supported Remsen & Graves' propositions, and suggested further that 
many sister taxa should be ranked specifically. 

Garcia-Moreno & Fjeldsa's analysis concentrated on southern Andean taxa. 
Their suggestion that the various Bolivian, Peruvian and some ^cudidox'mn Atlapetes 
be reassigned to two polytypic and six monotypic species has been followed by 
most subsequent works (Rodner et al. 2000, Ridgely & Greenfield 2001, Salaman 
et al. 2001, Hilty 2003, Dickinson 2003, Remsen et al. 2006). The northern taxa of 
A. latinuchus, A. schistaceus, A. albofrenatus and A. melanocephalus were not 
studied in detail, but various taxa formerly considered part of ^. rufinucha, namely 
baroni, caucae, chugurensis, comptus, elaeoprorus, latinuchus, nigrifrons 
{=phelpsi: see below), simplex {=spodionotus) and spodionotus, were tentatively 
reassigned to A. latinuchus. Garcia-Moreno & Fjeldsa noted 'Our study did not 
include . . . various more richly coloured forms in the northern Andes . . . However, 
it is evident that the current sequence does not reflect natural groupings'. 



Thomas M. Donegan & Blanca Huertas 107 Bull. B.O.C. 2006 126(2) 

Due to uncertainties in species limits amongst northern Atlapetes taxa and in 
order to describe A. I. yariguierum appropriately, we undertook a preliminary 
phylogenetic analysis of relevant taxa using a matrix of 16 adult plumage 
characters. We studied Garcia-Moreno & Fjeldsa's 'southern branch', 'central 
branch' and 'northern branch' taxa (except A. rufigenis) together with all other A. 
latinuchus, A. schistaceus and A. tricolor taxa, as well as A. fuscoolivaceus, A. 
albofrenatus and A. melanocephalus (which are considered closely related to 
'northern branch' species by Paynter 1978), A. I. yariguierum and a morphologically 
distinctive Atlapetes recently collected by A. Camero and A. Rodriguez in the Sierra 
de Perija ('Perija bird'). Chestnut-capped Brush-fmch Buarremon brunneinucha 
was used as an outgroup to permit comparison with Garcia-Moreno & Fjeldsa's 
phylogeny. Our matrix of character states was compiled from the literature (Paynter 
1978, Hilty & Brown 1986, Ridgely & Tudor 1989, Fjeldsa & Krabbe 1990, Ridgely 
& Greenfield 2001, Hilty 2003) and refined by examining specimens of all taxa 
(except A. latinuchus chugurensis and A. schistaceus taczanowskii). The characters 
and character states appear in Appendix 3. Separately, a preliminary phylogeny of 
northern Atlapetes taxa is being constructed by J. Klicka, C. D. Cadena & J. L. 
Perez-Eman through analyses of mtDNA genes cytochrome b and ND2. A sample 
of the A. I. yariguierum holotype is being analysed as part of this study and the 
preliminary data have been made available to us. 

In the morphological study, we first analysed taxa in Garcia-Moreno & Fjeldsa's 
molecular phylogeny for comparative purposes, using PAUP* v. 4.0 (Swofford 
2002), with characters unordered and equally weighted, with no re-weighting. As 
found by Garcia-Moreno & Fjeldsa, in our strict consensus tree close relationships 
were observed between: (i) Black-faced Brush-finch A. melanolaemus and Cuzco 
(Sooty) Brush-finch A. canigenis, (ii) Yellow-breasted Brush-finches A. latinuchus 
spodionotus and^i. /. comptus, and (iii) Slaty Brush-finches^, s. schistaceus and A. 
s. taczanowskii, in each case with high (>50%) bootstrap values. Conversely, 
Paynter (1978) hypothesised no close relationship between^, melanolaemus and^. 
canigenis. However, anomalously, A. r rufinucha was placed in our phylogeny as 
more closely related to A. latinuchus and A. schistaceus taxa, and A. tricolor as less 
closely related, than Garcia-Moreno & Fjeldsa found. We recognise the limitations 
of a study involving so few characters (see, e.g., Kitching et al. 1999), especially in 
a group in which pigmentation can be phylogenetically less informative than in 
other groups. However, an analysis, encompassing a broad range of plumage 
characters (cf. Paynter 1978) can be useful, if interpreted appropriately and 
conservatively and will provide results comparable with future molecular studies. 

Our matrix for all taxa in Garcia-Moreno & Fjeldsa's 'northern branch', as well 
as other Colombian and Venezuelan taxa within A. albofrenatus, A. fuscoolivaceus, 
A. latinuchus, A. melanocephalus, A. schistaceus and A. tricolor were subject to the 
same analysis. Three principal multi-taxa clades, each rooted from the same node, 
were identified in the strict consensus tree as follows: (i) A. schistaceus, subspecies 
castaneifrons, fumidus, schistaceus, taczanowskii and tamae; (ii) A. latinuchus. 



Thomas M. Donegan & Blanca Huertas 108 Bull. B.O.C. 2006 126(2) 

subspecies baroni, caucae, chugurensis, comptus, elaeoprorus, latinuchus, simplex 
{=spodionotus), spodionotus and yariguierum\ and (iii) A. melanocephalus, A. 
latinuchus nigrifrons and the Perija bird. The latter grouping was supported by 
bootstrap (83%); the others were not. All A. fuscoolivaceus, A. tricolor and A. 
albofrenatus taxa were unresolved in our strict consensus tree at the same level. The 
tree is not reproduced here as we do not propose a phylogeny. 

The current^, latinuchus complex (except^. /. nigrifrons, see below), including 
A. I. yariguierum, formed a monophyletic group. Preliminary data from Klicka et 
al.'s molecular analysis also suggest that^. /. yariguierum is most closely related to 
other A. latinuchus taxa (J. Klicka et al. in litt. 2005). However, this is a tentative 
arrangement, especially given the lack of data for other northern taxa such as ^4. /. 
caucae. 

A lack of significant genetic divergence amongst various taxa within A. 
latinuchus (A. I. chugurensis, A. I. comptus and A. I. spodionotus) was noted by 
Garcia-Moreno & Fjeldsa (1999). We further reveal no significant intra-group 
differences in morphometries (Appendix 2). Apparent intermediates between A. I. 
comptus and A. I. spodionotus are known from southern Ecuador and northern Peru 
(Ridgely & Greenfield 2001, Vellinga et al. 2004), suggesting a close relation 
between these taxa. Some morphological characters which vary within^, latinuchus 
are probably unreliable for delimiting species (Remsen & Graves 1995a). Presence 
or absence of a speculum is one case, having been reported as variously present and 
absent within the same population (Vellinga et al. 2004), and an apparent intergrade 
A. s. tamae I A. s. schistaceus (MLS 7552) had a speculum on one wing but not the 
other. Likewise, reported A. I. comptus I A. I. spodionotus intermediates and the 
surprising placement of some A. pallidinucha taxa in the A. latinuchus clade in the 
molecular phylogeny suggest that a paler nape is also somewhat plastic. 

Our analyses point to a clear anomaly in the current sequence (the placement of 
A. I. nigrifrons within A. latinuchus). The position of this taxon, discussed further 
below, can confidently be dealt with at this time in the light of the morphological 
data presented herein. However, a more substantial reappraisal of the A. latinuchus 
complex would be premature in advance of Klicka et al.'s more comprehensive 
molecular analysis. We have therefore described A. I. yariguierum as a subspecies 
within the A. latinuchus complex. However, we strongly suspect that some taxa or 
groups of taxa within this complex represent species under the Biological Species 
Concept (Helbig et al. 2002) and that most of them represent species under a 
phylogenetic species concept (Cracraft 1983). 

Before turning to the case of A. I. nigrifrons, its taxonomy requires comment. A. 
I. nigrifrons was described by Phelps & Gilliard (1940), but the name was 
subsequently changed to A. I. phelpsi by Paynter (1970) as the subspecific epithet 
was preoccupied by A. torquatus nigrifrons. However, with A. torquatus now 
assigned to Buarremon (see Hackett 1993, Remsen & Graves 1995a,b, and followed 
by almost all recent authors), phelpsi becomes a junior synonym of nigrifrons 
(Dickinson 2004). 



Thomas M. Donegan & Blanca Huertas 109 Bull. B.O.C. 2006 126(2) 

The clade including A. I. nigrifrons, A. melanocephalus and the Perija bird was 
the best supported of our northern taxa phylogeny. A. melanocephalus and A. I. 
nigrifrons would therefore, provisionally, appear to be more closely related to one 
another than either is to any of the A. latinuchus taxa, a proposition supported by 
biogeographical and morphological evidence. A. I. nigrifrons and A. 
melanocephalus are both restricted to the northernmost Colombian and Venezuelan 
mountains: the Perija and Santa Marta ranges. Studies by Remsen & Graves (1995a) 
and Garcia-Moreno & Fjeldsa (1999) suggest that some geographically close but 
morphologically distinctive Atlapetes taxa are more closely related to one another 
than they are to superficially similar but more geographically distant taxa. Paynter 
(1978) drew attention to the morphological similarity of ^. /. nigrifrons (then A. I. 
phelpsi) and A. melanocephalus, which share a black forehead (with A. 
melanocephalus possessing an entirely black crown), black chin, lack of pale 
moustachial markings (the malar merging with the mask), distinctly greyish cheeks 
and a paler grey back, features not found in northern A. latinuchus taxa. A lineage 
extending from A. a. albofrenatus (green back; moustachial stripe; black forehead; 
red crown), through the Perija bird (green back; moustachial merging with malar; 
black forehead; red crown) and A. I. nigrifrons (grey back; moustachial merging 
with malar; black forehead; red crown) to A. melanocephalus (grey back; 
moustachial merging with malar; black forehead; black crown), although not 
supported by our phylogenetic analysis, wherein A. albofrenatus taxa behaved 
counter-intuitively, appears a more plausible hypothesis than any close relation 
between A. I. nigrifrons and northern A. latinuchus taxa. 

The assignment of ^. /. nigrifrons to the A. latinuchus species-group appears to 
be a clear example of the current sequence failing to reflect natural groupings, per 
Garcia-Moreno & Fjeldsa (1995). If not assigned species rank, a better placement 
for nigrifrons is in either A. melanocephalus or A. albofrenatus. Other possible 
approaches, lumping A. melanocephalus and/or A. albofrenatus within A. 
latinuchus, defy rationale. Significant morphological differences exist between 
northern^, latinuchus, A. I. nigrifrons, A. melanocephalus andv4. albofrenatus, and 
are considerably greater than those between various southern forms now treated 
specifically. Under a modernised Biological Species Concept (Helbig et al. 2002) 
and given almost certain paraphyly of ^. latinuchus, we consider thaty4. /. nigrifrons 
is better treated specifically. The status of this taxon under a phylogenetic species 
concept should also be beyond doubt. Perija Brush-fmch is an appropriate 
vernacular name for A. nigrifrons. 

Finally, it merits further consideration that^. albofrenatus was not monophyletic 
in our analysis. The close resemblance of ^4. a. albofrenatus to A. schistaceus was 
discussed previously (Remsen & Graves 1995a). In our majority rule and Adams 
consensus trees, A. a. albofrenatus grouped with the five A. schistaceus taxa and not 
A. a. meridae. The lack of a very close relationship between ^1. a. meridae and^. a. 
albofrenatus was further supported by morphometric data: A. a. albofrenatus 
averages larger in both wing- and tail-length than A. a. meridae (Appendix 2; cf. 



Thomas M. Donegan & Blanca Huertas 1 1 Bull. B. O. C. 2006 126(2) 

Paynter (1978) who considered that 'no difference in size is apparent' between these 
taxa). No contact zone is known between them, A. a. albofrenatus being restricted 
to the Eastern Andes of Colombia and A. a. meridae to the Merida Mountains of 
Venezuela. Despite morphological differences between them, we suspect that A. a. 
albofrenatus is indeed the closest extant relative of A. a. meridae (though the 
converse may not be true given the probable link between A. a. albofrenatus, the 
Perija bird and^. /. nigrifrons, and possible links between^, a. albofrenatus and A. 
schistaceus). A. a. albofrenatus and A. a. meridae are morphologically more 
dissimilar (both in plumage and biometrics) than other closely rQlated Atlapetes taxa 
now treated specifically (following Garcia-Moreno & Fjeldsa 1995). They surely 
represent species under a phylogenetic species concept and quite probably under the 
Helbig et al. (2002) interpretation of the Biological Species Concept. However, 
insufficient evidence exists to suggest that A. albofrenatus is paraphyletic, and an 
analysis of genetic and/or vocal data is lacking. 

Conservation 

A. I. yariguierum is endemic to the Colombian East Andes EBA (038: Stattersfield 
et al. 1998). It technically qualifies for lUCN Category D2 Vulnerable status due to 
its Area of Occupancy being less than 1 00 km^ and being certainly known from 
fewer than five localities. However, if confirmed at all of the additional localities 
mentioned above, its status would be Near Threatened. Extensive field work in the 
main Cordillera in dptos. Santander and (particularly) Boyaca during the past 50 
years have yielded just a handful of specimens. The taxon appears to be somewhat 
rare even in primary habitat. Despite specific searches, field work at the type 
locality produced just one capture and no observations, and, at Lepipuerto, just one 
sighting and no captures. At our paramo site on the east slope of the Yariguies range, 
it was more common, being observed daily. Like other members of the A. latinuchus 
group (Hilty 2003), the new taxon appears to tolerate secondary habitats and may 
be more numerous in these than in primary forest. 

Protected areas in the Eastern Andes (e.g. Parques Nacionales Naturales Pisbe, 
El Cocuy, Tama, Sumapaz, Chingaza, Los Picachos, Alto Fragua, Indi-Wasi and 
Catatumbo) are concentrated in the highest paramos or on the east slope, and all are 
subject to varying levels of deforestation and human occupancy. Serrania de los 
Yariguies, one of the last forest wildernesses within this Critical-rated EBA 
(Stattersfield et al. 1998), was for 24 years subject to a 'conservation plan'. 
Following our work and with the impetus of the Ministerio de Medio Ambiente, 
Corporacion Autonoma Regional de Santander (CAS), various mayoralties of the 
region and NGOs, the Serrania de los Yariguies National Park was finally declared 
on 16 May 2005. The new protected area should assist in conserving A. I. 
yariguierum and other threatened birds including Black Inca Coeligena prunellei, 
Odontophorus strophium, Macroagelaius subalaris and Saffron-headed Parrot 
Pionopsitta pyrilia (Donegan et al. 2005, Donegan & Huertas 2005, Huertas & 
Donegan 2006). 



Thomas M. Donegan & Blanca Huertas 1 1 1 Bull. B. O. C. 2006 126(2) 

Comments on brush-finch vernacular names 

Garcia-Moreno & Fjeldsa (1999) suggested the English names 'Northern Rufous- 
naped Brush-fmch' for A. latinuchus and 'Bolivian Rufous-naped Brush-fmch' for 
A. rufinucha. However, 'Northern Rufous-naped Brush-finch' scarcely befits A. 
latinuchus, given that at least two taxa in the group {A. I. baroni and A. I. 
chugurensis) have white or cream napes, and that this name incorrectly implies A. 
rufinucha to be its sister species. Further, rufous-naped is a translation of the Latin 
species epithet oi A. rufinucha, not A. latinuchus. The name 'Cloud-forest Brush- 
finch' was also recently proposed (Clements & Shany 2001), but A. latinuchus is 
one of many Atlapetes species found in such zones and^. /. baroni, A. I. caucae and 
A. I. chugurensis are not restricted to cloud forests. Given this, J. V. Remsen in 
Dickinson (2003) proposed a third name, 'Yellow-breasted Brush-finch', which is 
consistent with the plumage of all group members and distinguishes it from some 
Atlapetes, but not from many others, e.g., Dusky-headed Brush-finch A. 
fiiscoolivaceus. Yellow-headed Buxsh-fmoh A. flaviceps, Tricoloured Brush-finch^. 
tricolor or Moustached Brush-finch A. albofrenatus, and also has the potential to 
confuse with Yellow-throated Brush-finch A. gutturalis. 'Cloud-forest Brush-finch' 
and 'Northern Rufous-naped Brush-finch' were both rejected by the AOU South 
American Checklist Committee. None of the English names proposed to date is 
particularly satisfactory, reflecting the lack of unifying plumage characters for A. 
latinuchus and that the group almost certainly contains several biological and 
multiple phylogenefic species. 'Yellow-breasted Brush-finch', whilst not ideal, was 
recently adopted by the AOU South American Checklist Committee (Remsen et al. 
2006). Doubtless a new suite of vernacular names will become necessary when A. 
latinuchus is, almost inevitably, split. The English name proposed herein for A. I. 
yariguierum is proposed only as a name for this taxon and not to any larger grouping 
(e.g. with^. /. elaeoprorus and/or^. /. spodionotus). 

Acknowledgements 

We thank Elkin Briceno for his help in the field and the rest of the EBA Project and YARE Project teams, 
Jorge Avendano, John Arias, Martin Donegan, Cristobal Rios, Laura Rosado and Diana Villanueva, and 
local guides Jose Pinto, Alonso Macias and Hernando Figueroa. Hernando Guevara (Corporacion 
Autonoma Regional para la Defensa de la Meseta de Bucaramanga — CDMB), Alvaro Prada Prada, 
Armando Rodriguez and Hector Lamo (CAS), and the mayors of San Vicente de Chucuri, Galan and El 
Carmen provided the necessary permissions for field work (CAS resolution no. 832). Robert Prys- Jones, 
Mark Adams and Douglas Russell (NHM), Enrique Castillo and Fernando Forero (lAVH), Jose Gregorio 
Moreno Patino (UIS), Roque Casallas and Arturo Rodriguez (MLS), Ray Symonds (UMZC), F. Gary 
Stiles (ICN) and J. F. & C. Voisin (MNFTN) provided access to specimens. Robin Restall provided 
photographs and biometric data of A. I. nigrifrons specimens in COP, and Mary Hennen digital 
photographs of material at FMNH. Sylke Frahnert (Berlin), Frank Steinheimer and Gerald Mayr 
(Frankfurt) provided invaluable data and insights concerning specimens in Germany. N. Krabbe offered 
information on Atlapetes in the Warsaw museum. Thanks to the Biomap Project (especially Juan Carlos 
Verhelst and Paul Salaman) for specimen data. J. Klicka, CD. Cadena and J. L. Perez-Eman sequenced 
the holotype and proffered their preliminary results. Richard Ranft (British Library) produced some of 
the sonograms. Jorge Avendano, Carlos Daniel Cadena, Roberto Chavarra, Oswaldo Cortes, Maria 
Angela Echeverry, Julian Idrobo, Diego Armando Rincon, Jorge Parra, Mike Sharpe, Clare Turner and 



Thomas M. Donegan & Blanca Huertas 1 12 Bull. B.O.C. 2006 126(2) 

Johana Edith Zuluaga informed us of their records of A. latinuchus. Alejandro Camero and Arturo 
Rodriguez provided data on the Perija bird. Thanks also to Ana Papadopolou (NHM). The expeditions to 
Serrania de los Yariguies were made possible due to the generous financial support of the Royal 
Geographical Society (with Rio Tinto pic), Duke of Edinburgh, Fondo para Accion Ambiental, 
Fundacion Omacha, Conservation International Colombia (Becas Iniciativa de Especies Amenazadas — 
Jorge Ignacio "El Mono" Hemandez-Camacho), the Percy Sladen Memorial Fund (Linnean Society) and 
Fundacion ProAves. YARE Project was supported by the BP Conservation Programme (BirdLife 
International, Conservation International, Flora & Fauna International, Wildlife Conservation Society), 
Game Conservancy Trust, Fundacion ProAves, World Pheasant Association, Universidad Industrial de 
Santander, Universidad de Caldas and Gobemacion de Santander. We thank Carlos Daniel Cadena, Jon 
Fjeldsa, Guy Kirwan, Niels Krabbe, J. Van Remsen and Paul Salaman for comments on the manuscript. 

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Addresses: Thomas M. Donegan, Fundacion ProAves, 33 Blenheim Road, Caversham, Reading RG4 
7RT, UK, e-mail: tdonegan@proaves.org / thomasdonegan@yahoo.co.uk. Blanca Huertas, 
University College London / Natural History Museum, London, UK, e-mail: b.huertas@nhm.ac.uk 
/ blancahuertas@yahoo.com 

APPENDIX 1 

Specimens examined. 

A. fuscoolivaceus: lAVH 7870, 11782, 11801; ICN 3169, 27333, 27336, 27370. 

A. melanocephalus: NHM 1885.6.8.125, 1885.6.8.452; lAVH 558, 944, 966, 2056-2058, 2064, 2067, 
2240,3085, 11300, 11311; ICN 23504-23515, 21690; MLS 7505 

A. latinuchus barontmiMmmxxmhQrQd, 1896.10.6.228, 1896.10.6.229 

A. latinuchus caucae ICN 3179, 3350, 19814, 25661, 25934, 25935, 26069, 29016, 29748, 29758, 
29804, ; MLS 7530, unnumbered; MNHN 2374 (NB: only Colombian^, latinuchus group speci- 
mens inspected in detail at MNHN). 

A. latinuchus comptus NHM 1883.6.12.930, 1885.6.8.149. 

A. latinuchus elaeoprorus NHM 1885.6.8.150, 1885.6.12.931; lAVH 2263, 2310, 11699; ICN 20169, 
34701; MLS 7528, 7530, 7870; MNHN 2375, 3754. 

A. I. latinuchus ^HM 1885.6.12.926, 1896.10.6.230-1896.10.10.233, 1900.10.2.93, 1924.2.14.22, 
1924.2.14.23, 1953.68.464. 

A. [latinuchus] nigrifrons ICN 32692; COP (photo) 55881, 55882, 55890, 55900, 55902, 55905, 
58251, 73053, 73323, 73326, 73333, 73337; ICN 32646 ('Perija bird'). 

'A. latinuchus simplex' (^ A. I. spodionotus) NHM 1893.12.12.18. 

A. latinuchus spodionotus ^YiM 1860.11.9.9, 1860.11.20.83, 1885.6.147, 1885.6.8.148, 1885.6.12.928, 
1885.6.12.929, 1900.10.2.87, 1900.10.2.88, 1925.12.24.283, 1925.12.24.284, 1938.12.20.122, 
1940.12.5.117-1940.12.5.120, 1940.12.5.214, 1940.12.5.960, 1940.12.5.961, 1969.52.520, 
1969.52.521, 1977.5.7-1977.5.11; UMZC 27/Fri(E)/8/r/l-27/Fri(E)/8/r/4; ICN-UN 33355; MLS 
7531,7532, MNHN 2376. 

A. latinuchus yariguierum ICN 3199, 10322, 25111, 34016 (holotype); UIS 1412; FMNH (photo) 
220606, 220607. 

A. a. albofrenatus ^HM 1885.6.8.152, 1885.6.8.153, 1885.6.12.932, 1885.6.12.933, 1916.9.21.141; 
lAVH 612, 6543, 10301, 11678, 12551; MLS 7533-7536; ICN 3171-3177, 3560, 3701, 3706, 
3721, 3974, 4217, 4819, 4820, 5068, 10323, 14712, 15938, 16250, 16358, 18847, 25280, 25497, 
28356, 28357, 18846, 18848, 31126, 34982; UIS 1433. 

A. albofrenatus meridael^UM 90.5.15 A, 1914.11.26.673-1914.11.26.676, 1914.11.26.680, 1969.39.77, 
1969.39.76, 1969.52.519. 

A. schistaceus castaneifrons ^YiM 12.18.070, 12.204.94, 85.6.8.160, 85.6.8.161, 85.6.12.941, 
1914.11.26.484, 1914.11.26.67, 1914.11.26.679, 1914.41.26.680-1914.11.26.683, 
1914.11.26.685-1914.11.26.687, 1915.3.1.177, 1915.3.1.678, 1969.39.78-1969.39.80. 

A. schistaceus fumidus ICN 4851, 4852. 

A. s. schistaceus ^WM 1845.5.24.30, 1854.1.25.100, 1857.11.28.59, 1885.6.8.156-1885.6.8.159, 
1885.6.8.162, 1885.6.12.934, 1885.6.12.935, 1885.6.12.938, 1898.12.14.654-98.12.14.657, 
1916.9.21.94, 1916.9.21.95, 1946.49.696, 2002.3.1044-1946.49.698; UMZC 
27/Fri(E)/8/5/l-27/Fri(E)/8/5/3; lAVH unnumbered (2), 5387, 6714, 7257, 7260, 7267, 11845, 
12287, 12627, 12646; MLS 7546-7548, 7550, 7551, 7553, 8245; ICN 4296^302, 4304-4306, 
4840, 4849, 4855-4864, 4866-4869, 4871-4884, 14655, 19015, 20010-20013, 20446, 
22330-22332, 26220, 26234, 33709, 35012, 437125. 

A. schistaceus tamae lAVH 10628, 10632, 10643, 10652, 12102, 12105; MLS 7537-7545; ICN 4853, 
4854, 10681, 33936; UIS 1252. 

A. canigenis NHM 1939.1.30.1. 



Thomas M. Donegan & Blanca Huertas 



115 



Bull. B.O.C. 2006 126(2) 



A. r. rufinucha-HmA 1846.9.9.99, 1846.99.136, 1885.4.8.144, 1885.6.8.142, 1885.6.8.143, 

1885.6.8.148, 85.6.12.922-1885.6.12.924, 1902.313.334. 
A. melanolaemus^mA 1902.3.13.335-1902.3.13.339. 

APPENDIX 2 

Biometrics oi Atlapetes taxa. 



Taxon 


Sex 


Wing 


Tail 


Tarsus 


Bill 


Mass 


A. fuscoolivaceus 


all 


74.0 ±3.4 (7) 


78.4 ±4.9 (7) 


26.6 ± 0.7 (6) 


16.4 ±0.6 (5) 


32.5 ±1.5 (2) 




males 


75.6 ±2.1 (5) 


80.0 ±5.5 (5) 


26.8 ± 0.9 (4) 


16.7 ±0.8 (3) 


32.5 ±1.5 (2) 




females 


67.0 ±0.0(1) 


73.0 ±0.0(1) 


26.0 ±0.0(1) 


16.0 ±0.0(1) 




A. melanocephalus 


all 


72.2 ± 3.5 (26) 


74.0 ± 3.2 (24) 


26.1 ±1.0 (27) 


16.0 ±0.6 (24) 


25.0 ±2.8 (6) 




males 


72.8 ±4.5 (10) 


76.0 ±3.5 (10) 


26.2 ±1.1 (10) 


15.9 ±0.5 (8) 


23.8 ±2.5 (2) 




females 


71.3 ±2.4 (12) 


72.2 ±2.2 (11) 


26.1 ±0.8 (12) 


16.0 ±0.7 (12) 


25.8 ±3.75 (3) 


A. latinuchus baroni 


all 


73.0 ±0.0 (3) 


76.7 ±2.5 (3) 


28.0 ±0.5 (3) 


15.5 ±0.0 (3) 






males 


73.0 ±0.0(1) 


79.0 ±0.0(1) 


28.5 ±0.0(1) 


15.5 ±0.0(1) 






females 


73.0 ±0.0 (2) 


75.5 ±2.1 (2) 


27.8 ± 04 (2) 


15.5 ±0.0 (2) 




A. latinuchus caucae 


all 


71.5 ±2.1 (12) 


72.3 ±2.7 (12) 


26.4±1.1(12) 


15.3 ±0.8 (11) 






males 


72.2 ± 2.4 (5) 


73.6 ±2.7 (5) 


26.2 ±1.5 (5) 


154 ±0.9 (5) 






females 


71.0 ±1.9 (7) 


71.3 ±24 (7) 


26.5 ± 0.8 (7) 


15.2 ±0.7 (6) 




A. latinuchus comptus 


all 


74.0 ±1.4 (2) 


74.5 ± 0.7 (2) 


27.5 ±0.7 (2) 


15.5 ±0.0 (2) 




A. latinuchus elaeoprorus 


all 


75.9 ±3.7 (10) 


73.7 ±44 (11) 


27.1 ±0.7 (11) 


15.9 ±0.5 (12) 


27.5 ±0.0(1) 




males 


75.5 ±3.5 (2) 


74.5 ±2.1 (2) 


27.0 ±0.0 (3) 


16.2 ±0.3 (3) 


27.5 ±0.0(1) 




females 


72.5 ± 4.9 (2) 


69.0 ± 2.8 (2) 


26.5 ± 0.7 (2) 


16.0 ±0.7 (2) 




A. I. latinuchus 


all 


72.6 ±3.5 (9) 


71.1 ±34 (9) 


26.0 ± 0.9 (8) 


15.6 ±0.4 (8) 






males 


73.8 ±4.1 (5) 


73.2 ±2.8 (5) 


26.3 ±1.0 (5) 


15.6 ±0.5 (4) 






females 


71.0 ±2.0 (3) 


68.5 ±2.1 (3) 


25.5 ±0.5 (3) 


15.6 ±0.3 (4) 




A. latinuchus nigrifrons 


all 
males 


69.6 ±3.3 (15) 
70.3 ±4. 1(6) 


71.0 ±3.6 (15) 
73.7 ±3.8 (6) 


25.5 ±0.0(1) 


15.9 ±0.5 (3) 


31.0±0.0(1) 




females 


68.9 ±2.7 (9) 


69.7 ±2.2 (9) 


25.5 ±0.0(1) 


16.0 ±0.0(1) 




'Perija bird' 


all 


72.0 ±14(2) 


76.3 ±1.8 (2) 


26.0 ±0.0(1) 


15.3 ±1.0 (2) 


28.5 ±0.7(2) 




females 


73.0 ±0.0(1) 


77.5 ±0.0(1) 


26.0 ±0.0(1) 


16.0 ±0.0(1) 


29.0 ±0.0(1) 


A. latinuchus simplex'' 


all (=male) 


78.0 ±0.0(1) 


77.0 ±0.0(1) 


27.0 ±0.0(1) 


(broken) 




A. latinuchus spodionotus 


all 


75.9 ±34 (29) 


75.2 ±5.1 (28) 


26.9 ±1.1 (28) 


15.7 ±0.5 (28) 






males 


76.4 ±34 (17) 


76.3 ±5.2 (17) 


26.7 ±1.1 (18) 


15.6 ±0.5 (18) 






females 


74.9 ±4.3 (7) 


73.9 ±5.7 (7) 


27.4 ±0.5 (6) 


16.0 ±04 (5) 




A. latinuchus yariguierum 


all 


75.3 ±2.1 (3) 


78.3 ±1.5 (3) 


27.0 ±1.7 (3) 


16.0 ±0.0 (3) 


22.6 ±0.0 (I) 




males 


77 ±0.0(1) 


80 ±0.0(1) 


25.0 ±0.0(1) 


16.0 ±0.0(1) 


22.6 ±0.0(1) 




females 


76 ±0.0(1) 




78± 0.0(1) 


28.0 ±0.0(1) 


16.0 ±0.0(1) 


A. a. albofrenatus 


all 


72.9 ±24 (36) 


73.9 ±3.5 (37) 


26.2 ±1.0 (36) 


15.9 ±0.5 (36) 


30.0 ±2.6 (6) 




males 


74.2 ±2.6 (17) 


75.2 ±2.9 (18) 


26.1 ±1.1 (18) 


16.0 ±0.6 (18) 


29.0 ±1.2 (4) 




females 


71.8 ±1.6 (10) 


72.6 ±3.0 (10) 


25.8 ±0.8 (9) 


15.9 ±0.5 (10) 


32.0 ± 4.2 (2) 


A. albofrenatus meridae 


all 


67.2 ±2.4 (9) 


68.0 ±2.9 (9) 


26.5 ±1.2 (8) 


15.7 ±0.7 (9) 






males 


68.0 ±1.7 (3) 


69.0 ±1.0 (3) 


26.7 ±0.8 (3) 


15.7 ±0.3 (3) 






females 


63.0 ±0.0(1) 


65 ±0.0(1) 


25.0 ±0.0(1) 


16.5 ±0.0(1) 




A. schistaceus castaneifrons all 


70.5 ±3.0 (18) 


73.0 ±4.2 (18) 


27.3 ±1.0 (18) 


15.7 ±0.5 (17) 






males 


70.9 ±3.0 (13) 


73.7 ±4.3 (13) 


27.4 ±0.8 (13) 


15.8 ±0.5 (12) 






females 


69.3 ±4.0 (3) 


71.3 ±3.1 (3) 


27.9 ±0.8 (3) 


15.7 ±0.8 (3) 




A. schistaceus fumidus 


all 


72.5 ± 3.5 (2) 


72.5 ±2.1 (2) 


27.3 ± 04 (2) 


15.8 ±0.4 (2) 






males 


75.0 ±0.0(1) 


74.0 ±0.0(1) 


27.5 ±0.0(1) 


16.0 ±0.0(1) 






females 


70.0 ±0.0(1) 


71.0 ±0.0(1) 


27.0 ±0.0(1) 


15.5 ±0.0(1) 




A. s. schistaceus 


all 


75.5 ±3.3 (81) 


77.5 ±4.5 (82) 


26.8 ±0.9 (83) 


15.0 ±0.6 (81) 28.2 ±2.6 (11) 




males 


78.0 ±2.2 (26) 


80.3 ±3.0 (25) 


27.0 ±0.7 (25) 


15.0 ±0.5 (23) 


28.5 ±1.3 (4) 




females 


74.3 ± 2.6 (28) 


764 ± 3.4 (27) 


26.7 ± 0.8 (29) 


15.2 ±0.7 (30) 




A. s. schistaceus 


all 


76.1 ±3.1 (46) 


78.7 ±3.7 (45) 


26.8 ± 0.8 (48) 


15.1 ±0.5 (46) 


28.0 ±3.0 (8) 


Eastern Andes 


males 


78.0 ± 2.2 (20) 


80.2 ±3.2 (19) 


27.0 ± 0.8 (20) 


15.0 ±0.5 (18) 


28.0 ±1.0 (3) 



Thomas M. Donegan & Blanca Huertas 



116 



Bull.B.O.C. 2006 126(2) 





females 


74.6 ±2.4 (19) 


76.7 ±2.9 (18) 


26.5 ± 0.6 (20) 


15.2 ±0.7 (21) 


27.8 ±5.5 (3) 


A. s. schistaceus 


all 


77.0 ±2.5 (12) 


79.4 ±3. 1(12) 


26.7 ±0.9 (10) 


15.2 ±0.7 (10) 


28.8 ±1.0 (3) 


Western and Central Andes males 


78.6 ±2.5 (5) 


81.3 ±2.9 (5) 


27.0 ± 0.4 (4) 


15.1 ±0.8 (4) 


30.0 ±0.0(1) 




females 


75.0 ±2.0 (3) 


78.7 ±3.5 (3) 


25.8 ±1.2 (3) 


15.0 ±1.0 (3) 




A. schistaceus tamae 


all 


74.5 ±2.9 (17) 


74.8 ±3.0 (18) 


27.4 ±0.8 (18) 


15.7 ±0.6 (15) 


26.4 ±1.5 (7) 




males 


75.9 ± 2.0 (9) 


76.0 ±2.0 (10) 


27.4 ±0.6 (10) 


15.8 ±0.5 (9) 


27.3 ±1.5 (4) 




females 


73.8 ±4.2 (4) 


72.8 ±4.6 (4) 


27.5 ±1.1 (4) 


15.5 ±0.5 (3) 


26.0 ±0.0(1) 


A. canigenis 


unknown 


69.0 ±0.0(1) 


68.0 ±0.0(1) 


27.5 ±0.0(1) 


17.0 ±0.0(1) 




A. r. nifinucha 


all 


69.1 ±4.1 (10) 


64.6 ±4.4 (10) 


25.9 ±1.0 (10) 


15.0 ±0.7 (10) 






females 


68 ±0.0(1) 


62 ±0.0(1) 


25 ±0.0(1) 


14.0 ±0.0(1) 




A. melanolaemus 


all 


68.8 ±2.9 (6) 


67.8 ± 3.8 (6) 


27.2 ± 0.5 (6) 


15.0 ±0.7 (6) 






males 


71.5 ±2.1 (2) 


71.0 ±4.2 (2) 


27.3 ± 0.4 (2) 


15.0 ±1.4 (2) 






females 


67.0 ±2.6 (3) 


65.7 ±3.1 (3) 


27.2 ± 0.8 (3) 


15.0 ±0.5 (3) 





APPENDIX 3 

Matrix of characters and description of characters used in phylogenetic analysis 



Taxon | Character 




2 


3 


4 


5 


6 


7 8 


9 


10 


11 


12 


13 


14 


15 16 


B. brunneinucha (outgroup) 










2 








1 2 





4 





1 


1 


1 


1 


A. fuscoolivaceus 







2 


1 








1 





4 














1 1 


A. t. tricolor 







1 


2 








1 





4 














1 1 


A. tricolor crassus 







1 


2 








1 





4 








1 





1 1 


A. melanocephalus 







2 











2 


1 





2 











2 


Perija bird 










2 








2 








1 














A. latinuchus baroni 




1 













1 





2 














1 


A. latinuchus caucae 













1 




1 





3 














1 


A. latinuchus chugurensis 




1 













1 





3 














1 


A. latinuchus comptus 




1 













1 





2 














I 


A. latinuchus elaeoprorus 













1 




1 





3 














1 


A. I. latinuchus 













1 




1 





2 














1 


A. latinuchus nigrifrons 


















2 








1 














A. latinuchus simplex 


















1 





3 














1 


A. latinuchus spodionotus 


















1 





3 














1 


A. latinuchus yariguierum 



















1 





3 














1 


A. a. albofrenatus 










2 








1 1 




















1 1 


A. albofrenatus meridae 










2 








1 2 





9 














1 


A. schistaceus castaneifrons 

















1 


1 














9 







A. schistaceus fumidus 

















1 


1 





















A. s. schistaceus 














1 


2 


1 1 





















A. schistaceus taczanowskii 

















9 
/• 


1 1 





















A. schistaceus tamae 




















1 1 





















A. canigenis 




















1 1 


1 


7 













1 


A. r carrikeri 


1 








1 








1 1 





1 













1 


A. r rujinucha 


1 














2 


1 1 





1 













1 


A. melanolaemus 


1 

















1 


1 


7 













2 



Matrix of plumage characters. 1. Belly (0=grey, l=yellow, 2=white); 2. Pale nape (0=absence, l=presence); 3. Crown 
(O=rufous, l=tawny/yellowish, 2=neither rufous nor tawny); 4. Olivaceous on back (O=none, l=tinge, 2=deep olive 
/ green); 5. Visible speculum (0=absence, l=presence); 6. Supraloral spot (O=none/vestigial, l=small, 2=large); 7. 
Chin (0=black, l=as breast or whiter); 8. Forehead (O=rufous/tawny, l=narrow black, 2=broad black); 9. Mottling 
on breast (0=absence, l=presence); 10. Dark malar (0=thick, l=strong, 2=moderate, 3=faint, 4=none); 11. Cheeks 
(0=black/as mask, l=dark grey, 2=pale grey); 12. Breast-band (0=absence, l=presence); 13. Melanism of flanks 
(0=light, l=strong); 14. Orange supercilium (0=absence, l=presence); 15. Throat and upper breast (O=homogeneous 
with belly l=distinctly lighter, 2=distinctly darker); 16. Paler moustachial region (0=absent, l=present). 

© British Ornithologists' Club 2006 



Jan-Hendrik Becking 1 1 7 Bull. B. O. C. 2006 126(2) 

Notes on Waterfall Swift Hydrochous gigas: 
I. Occurrence and nesting 

by Jan-Hendrik Becking 

Received 30 April 2005 

Waterfall Swift Hydrochous gigas (originally Giant Swiftlet Collocalia gigas) was 
described from a female taken by A. L. Butler, in 1900, at Semangkok (=Gap) Pass, 
at C.820 m, in Selangor, central Peninsular Malaysia (03°42'N, 101°45'E) (Hartert 
& Butler 1901). On 2 October 1900, M. E. G Bartels collected another specimen, at 
Pasir Datar (06''46'S, 106°56'E), on the south-west slope of Mt Pangrango, western 
Java (Finsch 1901, 1902, Bartels 1906), where it was not particularly rare (Bartels 
1915). Its distribution on Java is apparently restricted to the western part, with no 
sight records from central or eastern Java. The easternmost locality where it has 
been observed is the Kole Beres tea estate (c.07°12'S, 107°17'E), on the southern 
slope of Mt Patuha (Bartels 1931). For Sumatra, the first records were two 
specimens (male and female) taken by E. Jacobson in the Solok Mountains, Padang 
Highlands (c.00°48'S, 100°39'E), in July 1914 (Robinson & Kloss 1924). These are 
the only specimens from Sumatra, although there are sight records from Mt Kerenci 
(c.0r47'S, 10ri3'E), Mt Leuser National Reserve (c.03M5'S, 97°10'E) and 
elsewhere (Van Marie & Voous 1988). In Borneo, H. gigas is known only from sight 
records, some insufficiently documented (Smythies 1999). 

The species' biology is poorly understood, its behaviour unstudied, and no 
accurate population counts at individual colonies available. It is considered Near 
Threatened due to the probably small population and apparent disappearance from 
certain localities (Chantler 1999, Chantler & Driessens 2000, BirdLife International 
2001). I studied the Waterfall Swift in the field, at its breeding sites, paying special 
attention to behaviour and breeding. The present paper describes the nesting habits 
ofthe Waterfall Swift. 

My original aim was to monitor a nesting site at the foot of Mt Gagak, near the 
Mt Halimun complex in western Java, sited in a cave below a riverbed, with three 
large boulders forming a 'roof and a small waterfall at its front, in the River 
Cisekati (Becking 1971). The site was first discovered by the Bartels family in 1922 
and monitored by them until c.1939. It was no longer used in 1971, when I revisited 
the site, probably due to heavy deforestation in the environs. In October-December 
1971 and 1977 I searched for other nesting sites in western Java, along a number of 
montane rivers deemed suitable for the species. 

Nesting sites 

A total of seven nesting localities was discovered on three mountains in western 
Java: on Mt Pangrango-Gede (3,019 m), Mt Salak (2,21 1 m) and in the Mt Halimun 
complex (1,370-1,929 m). The species appeared widespread on these mountains, 



Jan-Hendrik Becking 



118 



Bull. B.O.C. 2006 126(2) 



especially where their specific biotope 
requirements were available. Nearly 
all nesting sites were near waterfalls or 
along rivers close by, in areas of 
submontane or montane, usually 
primary, forest far from any human 
activity. 

At a locality with nesting swifts, 
two types of nest sites were usually 
distinguished. Nests of the first or 
main type, c.80-90% of the total 
number observed, were at the waterfall 
itself The fall was usually in a rather 
open situation, often within primary 
forest, and large water masses fell 
steeply from a rock outcrop, producing 
a fall of considerable height. In such 
cases, nests were placed in small 
niches within the rock walls near the 
top of the fall, close to or even behind 
the water 'curtain', or beside of the fall 
(Fig. 1). Usually, nests at each site 
numbered 5-8, rarely more. 

The second type, numbering 
c. 10-20% of the total number of nests, 
were in nearby narrow gorges or 
ravines, where rapid streams or 

rivulets formed small cascades of water. The nests were beside the same river as the 
waterfall or on a nearby tributary, some just 20-30 m from the main waterfall site, 
and were placed in holes or niches in the vertical earth or rock wall, one to several 
metres above the stream and often 3-15 m apart. In such situations, nests were not 
directly exposed to water spray or seepage. Such nests were perhaps second-choice 
sites, and the number at each locality usually varied from one to four. At the well- 
studied Cicewol, Mt Salak site (see below), for instance, in 1977-78 there were two 
additional nests besides the c.7-8 nests at the fall: one 20 m downstream, and the 
other 10 m upstream, of the fall. 

I also found nests, singly or clumped 2-3, widely spaced and with no apparent 
association to any colony. These were sited along fast-running currents or rivulets 
within dense primary forest in the same regions. 




Figure 1 . Waterfall in the River Cicewol, on the south 
side of Mt Salak, western Java. The vertical rock wall 
beside the water 'curtain' contains Waterfall Swift 
Hydrochous gigas nests. Note the primitive ladder 
used to examine the nests (Jan-Hendrik Becking) 



Study site 

A nesting site below a relatively low waterfall (25 m) in the River Cicewol 
(06°46'S, 106°56'E), on the south side of Mt Salak at c.l, 100-1,200 m, was 



Jan-Hendrik Becking 



119 



BullB.O.C. 2006 126(2) 



^.rJ^- 





Figure 2. Nest of Waterfall Swift Hydroclwiis gigas containing an egg near the crest of Cicewol waterfall, 
Mt Salak. It is partially sited below the water curtain of the fall as can be noted from the vertical streams 
of water in front of the nest (Jan-Hendrik Becking) 



selected for more detailed observations. At this site, the river dropped from a 
vertical, east-facing rock surface, forming a waterfall with a pool below. The walls 
around the fall were clad in a short vegetation of ferns (particularly 
Hymenophyllaceae), hydrophilic dicotyledons (mainly Urticacea, Elatostema spp.), 
mosses (bryophytes) and liverworts. The site was slightly shaded by overhanging 
trees above and exposed to direct sunlight for only a few hours per day. 
Approximately 7-8 nests were sited near the top of the waterfall, below, behind or 
close to the water 'curtain', on ledges and in holes within the vertical rock and side 
walls of the fall. To closely examine the nests, bamboo ladders and scaffolds were 
constructed (Fig. 1), which were removed immediately following nest inspection to 
minimise disturbance. Construction, nest inspection and dismantling took c.2-3 
hours for 50% of the nests, with the other half examined the following day. Ambient 
temperature at the site was c.l5°C in the early morning, rising to 25°C at noon on 
sunny days with no cloud and falling to c.9-12°C at night. Relative humidity was 
close to saturation (96-100%). 



Jan-Hendrik Becking 



120 



Bull. B.O.C. 2006 126(2) 



Observations 



Seven nests were discovered at the 
Cicewol site on 30 October-5 
November 1977. They were located in 
the crescent-shaped andesitic rock 
surrounding the fall. More nests may 
have been present, as I subsequently 
found the remnants of a fresh egg in the 
pool, apparently from a nest not 
detected earlier. On 12 November I 
returned with my Indonesian assistants 
to closely examine the nests. Three 
nests near the top of the fall, close to or 
behind the water, were inaccessible and 
therefore not examined. Two of these 
possibly had young, from observations 
with binoculars. The other was too deep 
within a crevice to determine its 
contents. The four lower nests, at 
c.8-16 m, were examined: two each 
had a single egg at different stages of 
incubation, one fresh, the other hard- 
set; the third appeared to be under 
construction or repair, as fresh green 
moss and liverworts had recently been 
added to its rim; and the fourth nest 
held a very young nestling. Nest 
inspection was repeated on 20 and 27 
November, permitting description of 

the plumage development (see Part II). Heavy rainfall caused the water flow to 
approximately double by the 20th, with large tree stems and branches coming over 
the falls, and forcing me to abandon my camp site close to the river. 

In 1978 I returned to the same site in September-October. The same number of 
nests (7-8) was present. A notable degree of nest fidelity was found, as c.50% of the 
1977 nests were occupied. Although this visit was earlier in the season than in 1977, 
breeding had commenced and the nests were at different stages. Of the six lower 
nests examined, one had a single hard-set egg, one a very recently hatched nestling, 
three a small chick at various stages of development, and the last an older, feathered 
nestling. 

I continued to monitor this site subsequently, in 1982 (May-August), 1983 
(June-August), 1987 (September-November), 1989 (March- April), 1990 
(July- August), 1992 (May-July), 1994 (March-May), 1997 (April-May) and 2001 




Figure 3. Close-up of the same nest as in Fig.2 (Jan- 
Hendrik Becking) 



Jan-Hendrik Becking 



121 



Bull. B.O.C. 2006 126(2) 




Figure 4. Nest of Waterfall Swift Hydrochous gigas of the second type, i.e. not directly associated with 
a waterfall, but placed in the earth wall along the bed of a rapid rivulet, Mt Pangrango, western Java (Jan- 
Hendrik Becking) 

(July- August). During these visits — especially during the breeding season — I 
usually restricted myself to estimating the number of birds present and counting the 
nests visible from below. Prior to 1992 the nesting site was still intact and appeared 
to be occupied by the same number of pairs. By 1997, however, deforestation had 
commenced upstream nearby, and there was increased human activity. When I 
visited in 2001 I found the environment even more degraded by logging. Swifts 
were still present, albeit just c.2-3 pairs. Their continued presence indicates a high 
nest-site fidelity, as is known in other swift species. 



Nesting habitat and conservation status 

During the search for nesting sites of Waterfall Swift in 1977, the species appeared 
widespread on the three mountains investigated. Main nesting sites appeared to be 
sited at waterfalls in montane rivers within primary forest, whilst secondary nesting 
sites were in narrow river gorges. The preference for undisturbed forest areas was 
confirmed by developments at the Cicewol site, where degradation of the 
surrounding forest was coincident with a reduction in Waterfall Swift numbers. 
Given the rate of deforestation even in montane areas within the species' range and 
the swift's apparent fidelity to breeding sites, its conservation status is cause for 



Jan-Hendrik Becking 1 22 Bull. B. O. C. 2006 126(2) 

concern. A thorough investigation of its past and present occurrence may lead to its 
being classified as Vulnerable; see also BirdLife International (2001). At some 
colonies unscrupulous collectors may also be a threat. 

Acknowledgements 

I am greatly indebted to Dr Joost Brouwer, Lord Cranbrook and Dr David W. Snow for valuable 
suggestions and comments on an earlier draft. The study would have been impossible without the 
assistance of a semi-permanent crew of Sundanese-Indonesian helpers: Endang, Adi Nurhadi, Maman, 
Odik, Engkus, Adman and Ocang. They were indispensable and should share the outcome of this study. 
I am indebted to them also for their unceasing friendship and care, often under difficult conditions. In 
Sundanese I would say: 'Hatur Nuhun!'. Dr Charles T. Collins, as referee, made many usefiil comments 
on drafts of this paper. 

References: 

Bartels, E. 1 93 1 . Vogels van Kole Beres. Nat. Tijdschr. Ned.Indie 9 1 : 308-348. 

Bartels, M. 1906. Systematisch Ubersicht meiner Java-Vogel. J. Orn. 54: 497-519. 

Bartels, M. 1915. Zur Omis Java. Verhand. Orn.Gesellschaft Bayern 12: 199-200. 

Becking, J.-H. 1971. The breeding of Collocalia gigas. Ibis 113: 330-334. 

BirdLife International. 2001. Threatened birds of Asia: the BirdLife International Red Data Book. 

BirdLife International, Cambridge, UK. 
Chantler, P. 1999. Family Apodidade (swifts). Pp. 388^57 in del Hoyo, J., Elhott, A. & Sargatal, J. (eds.) 

Handbook of the birds of the world, vol. 5. Lynx Edicions, Barcelona. 
Chantler, P. & Driessens, G. 2000. Swifts: a guide to the swifts and treeswifts of the world. Second edn. 

Pica Press, Robertsbridge. 
Finsch, O. 1 90 1 . [A note on the occurrence of Collocalia gigas. Hart. & Butler, in Java.] Bull. Brit. Orn. CI. 

12: 30. 
Finsch, O. 1902. Zur Verbreitung von Collocalia gigas Hartert et Butler. Orn. Monatsber. 10: 38. 
Hartert, E. & Butler, A. L. 1901. [Collocalia gigas, Hartert & Butler, n. sp.] Bull. Brit. Orn. C/. 11: 65. 
Robinson, H. C. & Kloss, C. B. 1924. On a large collection of birds chiefly from West Sumatra made by 

Mr. E. Jacobson. J. Fed. Malay States Mus. Kuala Lumpur 11: 189-374. 
Smythies, B. E. 1999. The birds of Borneo. Fourth edn. Natural History Publications, Kota Kinabalu. 
Van Marie, J. G. & Voous, K. H. 1988. The birds of Sumatra: an annotated check-list. British 

Ornithologists' Union Checklist No. 10. British Ornithologists' Union, London. 

Address: Ericalaan 7, 6703 EM Wageningen, The Netherlands, e-mail: janhendrikbecking@hotmail.com 
© British Ornithologists' Club 2006 



Jan-Hendrik Becking 123 Bull. B.O.C. 2006 126(2) 

Notes on Waterfall Swift Hydrochous gigasi 

II. Nestling plumage and phylogenetic 

relationships 

by Jan-Hendrik Becking 

Received 30 April 2005 

Waterfall Swift Hydrochous gigas occurs in montane Peninsular Malaysia, Sumatra, 
Java and possibly Borneo (see Part I, pp. 117-122). Originally placed in Collocalia 
(Hartert & Butler 1901), its taxonomic position is, however, still debated. For a 
Collocalia, Waterfall Swift is atypically large. Brooke (1970) introduced 
Hydrochous as a subgenus name, merely on the basis of literature information 
(without seeing a specimen or live bird), but later raised the name to genus level, 
with Waterfall Swift Hydrochous gigas as its sole member (Brooke 1972). Brooke's 
rationale was (1) the larger size of Waterfall Swift compared to other Collocalia, (2) 
its lack of echolocation ability (Medway & Wells 1969) in contrast to Aerodramus, 
and (3) its peculiar nesting sites, near or behind waterfalls (Somadikarta 1968, 
Becking 1971). The species was tentatively placed by him in the tribe Collocaliini 
of the subfamily Apodinae, following the general division of the Collocaliini into 
three genera: (a) duU-plumaged echolocators (Aerodramus), (b) duU-plumaged non- 
echolocators (Hydrochous) and (c) dull or glossy plumaged non-echolocators 
(Collocalia sensu stricto) (Brooke 1970, 1972). This division needs revision, 
however, because Collocalia troglodytes of the Philippines has recently been proven 
to echolocate (Price et al. 2004). 

To obtain insight into its phylogenetic relationships, Lee et al. (1996) compared 
mitochondrial cytochrome-Z? DNA sequences of Hydrochous gigas with those of 
several related species. The Maximum Likelihood Tree and the constructed 
Bootstrap Consensus Tree indicated that H. gigas is probably closely related to 
Aerodramus, although its precise phylogenetic position is uncertain because most 
parsimonious trees placed Hydrochous variously within Aerodramus, but never as a 
sister taxon to that genus. Thomassen et al. (2003) repeated the analysis but 
screened for a larger section of mitochondrial cytochrome-Z? DNA (i.e. 1143 bp, 
rather than 406 bp) in H. gigas and several allies. Their resulting Bootstrap 
Consensus Tree placed Waterfall Swift between two Aerodramus species, A. 
maximus and A. fuciphagus, but closer to A. maximus. The Maximum Likelihood 
Tree revealed the same topology, except that H. gigas now grouped with the single 
A. vulcanorum specimen examined. 

Thus, the precise phylogenetic relationships of the Waterfall Swift remained 
somewhat uncertain. It is remarkable, however, that both research groups uncovered 
evidence of a close relationship between Hydrochous and the echolocating 
Aerodramus species: Hydrochous gigas has been shown to lack echolocation 
capacity by experimentally letting a specimen fly in a dark room (Medway & Wells 



Jan-Hendrik Becking 1 24 Bull. B. O. C. 2006 126(2) 

1969). In a recent publication, Price et al. (2005) performed another DNA analysis 
(cytochrome-Z) gene) of the same Hydrochous gigas specimen studied by Lee et al 
(1996), which was collected by me and was examined by all of these authors 
without my knowledge or consent. Price et al. (2005) considered Hydrochous to be 
sister to the Three-toed Papuan Swiftlet Aerodramus papuensis (see Somadikarta 
1967). Moreover they concluded that both Hydrochous and A. papuensis are sister to 
all other Aerodramus swiftlets. Hence, under this arrangement these two species are 
removed from amidst the Aerodramus and are placed as basal to them all. 

My aim here is to describe the morphology and nestling plumages of Waterfall 
Swift in their various stages, as these might shed some light on the phylogeny of the 
species. 

Methods 

Observations 

Most observations of Waterfall Swift nestlings were made at a site in western Java, 
at a relatively low waterfall (c.25 m) in the River Cicewol (06%6'S, 106°56'E), on 
the south slope of Mt Salak, at an altitude of c. 1,100-1, 200 m. Permanent 
observation of the nests at this site was impossible due to access difficulties and the 
problem of disturbance (see Part I). During brief nest inspections, the nestlings that 
could be reached were examined for size, the presence or absence of feathers 
(semiplumes) and development of feather tracts (pterylae). Weight was measured 
using a Pesola balance. Colour descriptions of the naked parts and feathers 
(semiplumes, etc.) were matched using Ridgway (1912) and Smithe (1974) colour 
swatches. Black-and-white and colour photographs were made for reference. Most 
observations were made in 1977-78, supplemented subsequently by further 
observations at the same site. 

Because of the laborious work of constructing the stagings to reach the nests, 
nest inspections were usually repeated only 2-3 times after the first examination. 
Re-inspection was usually at an interval of 7-10 days, weather conditions 
permitting. Less-accessible nests were generally only visited once. Age estimations 
were therefore rather approximate, especially as the age of a chick when first found 
was not precisely known. Despite such shortcomings and the rather restricted 
number of visits, the approximate ages of the chicks were estimated reasonably 
precisely by comparing different chicks. Where possible, they were checked in 
following years and supplemented with additional information from other nesting 
sites. 

Semiplume morphology 

The 'pseudo-down' (semiplume covering) of Waterfall Swift nestlings was 
compared with that of Common Swift ^/?t/>y apus. Feather structures were studied in 
detail at higher magnifications with the aid of a Wild M-5 stereomicroscope 
(enlargements 60-300x) and a Wild M-20 research microscope (300-600x), in 
unstained preparations in air, observed under cover glass. The semiplume of the 



Jan-Hendhk Becking 125 Bull. B.O.C. 2006 126(2) 

Common Swift nestling examined was from a bird found dead in a nestbox, at 
Bennekom, The Netherlands, containing two other, live nestlings. It weighed 14 g 
and had a wing-length of 38.3 mm, and was estimated to be c.l4 days old. 

Anatomical study 

Because the course of the main arteries (carotid arteries) in the neck and thorax of 
Apodiformes (Apodidae) is important for a judgment on their phylogeny, these were 
studied in several alcohol specimens of Hydrochous gigas (Java) and Collocalia 
linchi (Java), and in fresh (frozen) specimens of Apus apus (Netherlands). Attention 
was also paid to the feet of//, gigas, i.e. the grip, and to the number of phalanges in 
the different digits, compared to those in other swift species. 

Photographic records 

As it was difficult to photograph the nestlings in situ on primitive ladders or 
scaffolds, the nestlings were briefly removed from the nests and photographed in a 
empty nest of the species at the foot of the fall, and afterwards immediately returned 
to their nests. The recently hatched young featured in Fig. 1 was found dead in its 
nest at another colony. 

Results 

Post-hatching development 

Four different developmental stages of the nestling were recognised. 

Newly hatched chick 

As in all Apodidae, the hatchling is naked and blind (Figs. 1-2, two different 
chicks). A well-developed whitish egg tooth is conspicuous at the distal upper ridge 
of the upper mandible of the pinkish bill. The lower mandible protrudes somewhat 
beyond the upper mandible due to a second egg tooth-like structure or tubercle at 
the distal end of the lower mandible. This was pointed out to be a second egg tooth 
(C. T. CoUins in litt. 2005). The skin is pinkish (Smithe: Salmon Color/Pink [6/7]; 
Ridgway: Pale Flesh Color (PI. XIV)), with a very slight plumbeous or greyish tinge 
(Smithe: close to Light Neutral [85]; Ridgway: Light Varley's Gray [PI. XLIX]). 
This greyish tinge is darker on the head, back and wings. The pinkish feet are soft 
and rather large for the size of the chick (Fig. 1). Newly hatched chicks weighed 
c.2-3 g and had a body length of c.3.5 cm. A large throat pouch is visible, an 
adaptation for taking food boluses. At this stage, the parents brood the chick 
continuously, and leave it only reluctantly if disturbed. The naked chick clearly 
requires nearly continuous parental cover for its insulation. 

Chick of 4-8 days 

At this age chicks are still completely naked and their eyes closed, but on the back 
slightly darker plumbeous feather tracts are visible below the epidermis, one dorsal 
(spinal tract) and one somewhat laterally on each side (femoral tract) (Fig. 3). In 



Jan-Hendrik Becking 



126 



Bull. B.O.C. 2006 126(2) 




Figure 1 . Recently hatched Waterfall Swift Hydrochous gigas nestling (Jan-Hendrik Becking) 




Figure 2. A few days-old nestling of Waterfall Swift Hydrochous gigas (Jan-Hendrik Becking) 



Jan-Hendrik Becking 127 Bull B.O.C. 2006 126(2) 

addition, within the apteria many small dark spots appear (the underlying follicles 
of the down-like semiplumes). The coronal and occipital tracts of the body also 
become more prominent as dark grey (Smithe: Medium Neutral Gray, 84) areas or 
lines. The underside and anal region are still very pale greyish pink, and no 
underlying feather tracts are visible. Subsequently, the plumbeous grey back and 
head become darker (Smithe: Dark Neutral Gray/Medium Neutral Gray, 83/84), 
whilst the pinkish bill acquires a blackish tip and darker upper rim. The skin of the 
orbits of the protruding eyes also becomes darker, as does the skin of the tibia and 
femur and at the upper rim of the wing. The base of the broad bill, however, is pale 
flesh pinkish, even almost white. At this stage the skin does not cover the body 
smoothly but is rather wrinkled or folded, especially on the back and body-sides. 
The gape and palate are vivid flesh-pink, the tongue pinkish with a greyish tip. 
When 6-7 days old the chick is c.5 cm in length (head width 15.6 mm), but with its 
neck stretched — as when begging — can reach 6.5-7.0 cm. It weighs c.9.3 g, i.e. 
about 1)-A times its weight when newly hatched (2-3 g, N=5). 

Chick of 10-15 days 

A 'downy' semiplume covering sprouts on the back from the developed follicles 
visible earlier, giving the chick what looks like a coat of down (Fig. 4). The 
semiplumes sprout mainly from the apteria, but are often denser at its borders, and 
also develop on the head and upper chest. The chick starts to open its eyes, but if 
exposed to brighter sunlight immediately closes them. When handled the chick 
always tries to turn its head towards the shade. The growth of the covering is rather 
fast on the dorsal side, but far slower on the ventral side. The semiplumes are 
greyish black (Smithe: Dark Neutral Gray, near 83) and when fully grown measure 
c.lO mm. The first wing- and tail-feathers appear, still enclosed by their sheaths. 

Chick of c.17-22 days 

Although the semiplumes are still prominent, the growth of the primaries and 
secondaries, followed by the greater coverts, is rather rapid. As these feathers are 
still in sheath, the chick acquires a pin-cushion-like appearance, an image 
strengthened because the sheaths of the greater coverts and those of the carpal edge 
often point obliquely sideways from the wing (Fig. 5). Later, the secondaries and 
upperwing-coverts start to open their vanes at the tips, whilst the primaries are still 
enclosed within their sheaths. 

There are indications that near the end of the 'downy' semiplume stage these 
semiplumes are very loosely implanted in the skin. A chick of this age found 
recently dead in a nest, which I tried to preserve as a specimen, lost most of its 
semiplumes during preparation, although I took utmost care to prevent this. The 
semiplumes which became detached during the skinning process were not still 
ensheathed, but full grown. 

At a late stage the dense semiplumes become mixed with another type of unique 
feathers, which sprout from other follicles. They are more like normal contour 



Jan-Hendrik Becking 



128 



BullB.O.C. 2006 126(2) 




Figure 3. A 7-9-day-old nestling of Waterfall Swift Hydrochous gigas showing the somewhat contrasting 
darker pterylosis tracts regions on the back, and (in the upper left comer) the copious quantities of whitish 
gelatinous saliva used to bind the nest to the rock (Jan-Hendrik Becking) 




Figure 4. A c.IO-15-day-old nestling with semiplumes and its llrsl contour feathers, the latter still in an 
early stage within their sheaths (Jan-Hendrik Becking) 



Jan-Hendrik Becking 



129 



BullB.O.C. 2006 126(2) 



feathers in having a loose-webbed base, followed by a short closed vane and ending 
in a more or less semiplume-like apex (Fig. 6d). The plumulaceous bases (of these 
new feathers) bear blackish-grey barbs identical to the initial semiplumes, but the 
closed vane is buffish yellow (Smithe: Cream Color, 54) and the grey semiplume- 
like endings have fine yellowish tips, affording the chick a somewhat variegated 
appearance, which is intensified when it raises its back feathers and appears bristly 
(Fig. 5). 

A second wave of largely plumulaceous feathers arising from other follicles is 
exceptional and conflicts with any moult system known. Moreover, moult is 
described as a periodic shedding and replacement of feathers (Campbell & Lack 
1985), and the follicles should persist and produce a series of feathers from each 
during the course of the bird's lifetime. This would mean that the downy 
semiplumes are overgrown by incoming contour feathers and thus disappear from 
sight, but are not lost or dropped. However, when I removed the contour feathers 
from an adult alcohol specimen of H. gigas I found very few 'downy feathers' 
below. This is a puzzle that requires solution. A second set of semiplumes is 
certainly very exceptional and unknown for any species of swift, but has been 
reported in other birds. It is unknown or disputed whether the moult pattern of 
contour feathers in birds can be applied for the semiplumes occuring in the apteria. 



MTr-^^- 










Figure 5. A c. 1 7-20-day-old nestling showing the unique type of feathers (intermediate) between 
semiplumes and contour feathers. When these feathers are raised the nestling acquires a bristly 
appearance (Jan-Hendrik Becking) 



Jan-Hendrik Becking 130 Bull. B.O.C. 2006 126(2) 

It has been suggested that I had to examine the semiplume downy cover (and 
folhcles) in the Waterfall Swift chicks more precisely, but this can not be done in a 
living chick without doing it severe harm. This can only be done in a dead chick, 
but I refrained from to sacrifice a chick for this purpose as the scope of this 
investigation was to follow its development'. 

Nestling of c.25-45 days 

At this stage nestlings were rarely handled, because they were rather sensitive to 
disturbance. Moreover, they tended to grip with their feet very strongly to the 
underlying nest material and were therefore difficult to remove. The chicks at this 
stage are covered with sooty grey-brown contour feathers (Smithe: between 
Blackish Neutral Gray (82) and Dark Neutral Gray (83); Ridgway: Deep Slaty 
Brown, Plate L), paler on the underside and darker on the upperside, and with 
extensive white at their bases, particularly on the back, belly and flanks. Often this 
basal white is not completely concealed, giving the nestlings a somewhat spotted 
appearance. Completely white small underlying feathers can also be found on the 
back and belly, but apparently only very few semiplumes under these first contour 
feathers, as far as could be determined. It is noteworthy that Waterfall Swift 
nestlings at this age lack the pale greyish fringes to the primaries, secondaries and 
other contour feathers well known in older nestlings or juveniles ofApus, and also 
present in some Cypseloidinae (Marin & Stiles 1992). I estimate that nestlings leave 
the nest 48-55 days after hatching. 

Description of the semiplumes 

Unlike true natal down (neossoptiles), semiplumes are modified loose-webbed 
contour feathers (teleoptiles). Semiplumes have a definite rachis but no hamuli on 
the barbules, and therefore cannot produce a firm vane (Nitzsch 1840, Chandler 
1916, Van Tyne & Berger 1971). The semiplumes of Hydrochous gigas are blackish 
grey (Smithe: Medium Neutral Gray/Dark Neutral Gray, near 84/83) and thus 
distinctly greyer than the uniform sooty-black (Smithe: Blackish Neutral Gray, 82) 
semiplumes of Apus apus studied for comparison. 

As mentioned, there are two successive types of plumulaceous feathers. The first 
(Fig. 6a) develops from the naked chick. These are more or less spherical in form 
and usually c.6-10 mm long, occasionally 12.5 mm when fully grown. Smaller ones 
may also occur, 5-6 mm long. The semiplumes of //. gigas have no aftershaft at 
their base (Fig. 6a), unlike the semiplumes of Apus apus. At low magnification the 



' From the observation that there are very few semiplume feathers in an adult it may be concluded that 
they are are lost and not replaced by new ones. It may be also that the dermal papilla at the base of the 
follicle is dormant for a time. I have the impression that the same sequence of processes, from 'downy 
cover' to juvenile and adult plumage, occurs in Apus apus. Through the courtesy of the Apus Working 
Group Netherlands, I have been given some chicks (found dead in the nest) and semi-adults and adults 
from Bird Shelters (mainly traffic victims), and with this material I hope to solve this plumage problem. 



Jan-Hendrik Becking 



131 



Bull.B.O.C. 2006 126(2) 




Figure 6. Drawings of semiplumes, etc. of Waterfall Swift Hydrochous gigas: (a) 'downy feather' 
(semiplume) of the first type; (b) a semiplume of the first type at low magnification, showing a chain of 
regularly spaced black dots (nodes) within the clear barbules; (c) a barbule of a semiplume of the first 
type at higher magnification, showing small thickenings (nodes or barbicels), which usually bear distally 
oriented spiny projections (prongs); and (d) a feather of the other unique type, being intermediate 
between a semiplume of the first type and a contour feather; it has a plumulaceous base followed by a 
short closed vane and ends in a fine semiplume-like apex (del. Jan-Hendrik Becking) 



Jan-Hendrik Becking 132 Bull. B. O. C. 2006 126(2) 

barbules of the semiplumes of Waterfall Swift appear transparent with a chain of 
regularly spaced black dots like a string of beads (Fig. 6b), whereas the barbules of 
A. apus are uniformly blackish without hyaline interspaces. At higher magnification 
(500-600X), the black dots in the barbules proved to be dark pigmented thickenings 
or projections, so-called nodes (Rosalind & Grubh 1987, Rajaram 2002). These are 
regularly spaced along the barbules with an intemodal distance of c.9-10//m. They 
are broadest towards the tip and usually have spiny projections, or 'prongs' (Fig. 
6c). 

The second type of feathers, being more like contour feathers, possess a partially 
closed vane and are about twice as long as the first type, c. 18-19 mm, width 11-12 
mm (Fig. 6d). When both types are observed closely with a magnifying glass or 
stereomicroscope, they are seen to have yellowish tips, a feature absent in the 
semiplumes of Apus apus. 

Anatomical study 

The Cypseloidine swifts, which in breeding in association with waterfalls and also 
in their size, are very similar to that of Hydrochous gigas. These Cypseloidine swifts 
have, however, two carotid arteries (Glenny 1953, 1955), as in most birds, which is 
supposed to be the primitive conditon. In H. gigas I found only one carotid artery in 
the laevo, i.e. sinistra, position connected to the heart, as in Collocalia linchi and 
Apus apus. The latter is regarded as the derived condition. In contrast to the feet of 
representatives of the genus Apus (Stresemann 1934), those of H. gigas are 
anisodactyl without any reduction of phalanges in the toes. This means its feet have 
3, 4 and 5 phlanges for digits 2, 3 and 4, respectively. 

Egg tooth development 

The observed second egg tooth on the lower mandible of the Waterfall Swift is not 
unique to the species, having been found in other swifts. Collins & Naik (1975) 
described it for Apus nipalensis, of the subfamily Apodinae, in India. Moreover, it 
is reported (Collins 1968) for Short-tailed Swift Chaetura brachyura and Chestnut- 
collared Swift Cypseloides rutilus of the subfamily Cypseloidinae. Although of 
interest, the presence of two egg teeth in newly hatched H. gigas chicks throws no 
light on the phylogenetic relationships of Waterfall Swift. 

Breeding biology compared to other swift species 

Based on my observations, the breeding season of Waterfall Swift in western Java 
appears to last from September to January, sometimes until February/March (due to 
replacement clutches), coinciding with the rainy season, when termites, preferred 
food of Waterfall Swift, conduct mating flights and are readily available (pers. obs.). 
Termites were also found to be usually the sole prey in stomachs of this species 
examined by Max Bartels Snr. (notes held in the Leiden Museum). Moreover, the 
few stomachcs (five), which I examined (in the rainy season) were distinctly 
enlarged and crammed with remains of termites. The estimated fledging period of 



Jan-Hendrik Becking 1 33 Bull. B. O. C. 2006 126(2) 

45-55 days is slightly shorter than that of the similar-sized and ecologically similar 
White-chinned Swift Cypseloides cryptus of the Neotropics (Marin & Stiles 1992, 
Chantler 1999). There is also a great similarity between the breeding biology of 
Hydrochous gigas and Cypseloidine swifts of the New World in respect of site 
preference, nest type and, for some species, even clutch size (single-egg clutches). 
H. gigas and Cypseloidine swifts are not, however, closely related, as fundamental 
differences exist between them (see above). The similarity is doubtless due to 
convergent evolution, resulting in their adaptation to the same very special nesting 
requirements. 

Discussion 

The discovery of a 'down-like' semiplume nestling plumage in Hydrochous gigas 
nestlings is surprising, as it is absent in all species considered its closest relatives, 
i.e. Collocalia and the echolocating Aerodramus. It is also lacking in the tribe 
Chaeturini (following Brooke 1970). Such plumage is, however, present in the 
subfamily Cypseloidinae of the New World, which also shows a great resemblance 
to Waterfall Swift in morphology, habits and nest-site choice, selecting sites close to 
running water or waterfalls, and construct similar types of nest (Becking 1971, 
Marin & Stiles 1992). 

The semiplume covering of young nestlings of Hydrochous is probably an 
adaptation to its breeding in cool damp environments, assisting the insulation and 
thermoregulation of the chick, a view supported by some of my field observations. 
Newly hatched nestlings of Waterfall Swift were near-continuously brooded by one 
adult, but when the semiplume plumage was fully developed they were left 
unattended for shorter or longer periods. A similar suggestion regarding the function 
of down-like plumage was also made for Cypseloidine nestlings by Legg (1956) and 
Collins (1963). 

The formation of a second plumulaceous covering, as reported for H. gigas, 
appears to be unique in swifts, but a second, successive coat of nestling down was 
first noted by W. E. Clark (1906) in penguins (for neossoptiles), and later recorded 
in most owls as well as certain other groups. These, however, are from the same 
follicle and are extruded on the incoming contour feathers (teleoptiles). 

Environmental circumstances may not be the only explanation for the forming 
of a down-like covering. Of the two species of Cypsiurus palm swifts living under 
similar climatic conditions to each other, with the same life histories and nest-site 
choice, and building the same type of nests, the chicks of African Palm Swift C 
parvus are densely covered with semiplumes (Schuster 1912, Moreau 1941, Collins 
1965). Chicks of Asian Palm Swift C balasiensis, however, are completely naked 
until the contour feathers appear (Hails & Turner 1984; pers. obs.). This suggests the 
presence of an essential genetic factor, playing a role in the growth (i.e. presence or 
absence) of semiplumes on chicks, in addition to environmental factors already 
mentioned. 



Jan-Hendrik Becking 134 Bull. B.O.C. 2006 126(2) 

Teleoptile semiplume plumage is also present in chicks of Apus apus and Alpine 
Swift Tachymarptis melba, which were originally cliff breeders in a cool climate 
(Glutz von Blotzheim & Bauer 1980, Cramp 1985). Unlike Apus apus and 
Tachymarptis melba, however, which have a so-called pamprodactyl foot (but see 
Collins 1983) with only three phalanges in digits 2-4 (Stresemann 1934), those of 
H. gigas are anisodactylous with no reduction in the phalanges (see above). The 
subfamily Cypseloidinae similarly have the normal avian anisodactylous condition, 
as does the enigmatic African genus Schoutedenapus . For precisely this reason. 
Scarce Swift Schoutedenapus myoptilus underwent a generic name change from its 
original Apus myoptilis (Salvadori 1888, De Roo 1963). There is a chronic lack of 
available data for Schoutedenapus: its breeding is unknown, and little is known of 
its behaviour and distribution either. The ignigmatic position of both 
Schoutedenapus myoptilus and Hydrochous gigas was reviewed by Collins (2000). 

It is striking that the two Cypseloides with the same mass as Waterfall Swift 
(35.79 g, A/=19, pers. obs.), namely C. cryptus (35.27 g, 7V=13) and C niger (35.71 
g, 7V=16), also lay only a single egg (Legg 1956, Marin & Stiles 1992). The much 
smaller C. rutilus (21.32 g, N=\?>9) has a two-egg clutch (Collins 1968, Marin & 
Stiles 1992). There may be a relation between body mass and nesting behaviour in 
these species. A close relationship between Cypseloidine swifts and Waterfall Swift 
is unlikely, for several reasons. All Cypseloidine swifts have a diastataxic wing (H. 
L. Clark 1906, Stephan 1970) and two carotid arteries like most birds (Glenny 1953, 
1955), whereas some Apodini (Collocalia bartschi and Aewnautus andecolus) have 
only one laevo-carotid artery (Glenny 1953, 1955). As mentioned, I found only one 
carotid artery, in the laevo, i.e. sinistra, position, in Waterfall Swift, Collocalia 
linchi and Apus apus. Moreover, Cypseloidine swifts possess no active salivary 
glands and therefore do not use saliva for nest building (Johnston 1961, Marin & 
Stiles 1992, Marin 1997). Waterfall Swift, however, uses copious saliva for nest 
attachment and some to bind the nest material, especially the rim. This saliva is 
originally white or opaque whitish gelatinous (see Fig. 3), but soon becomes black, 
particularly in older nests and especially in museum specimens. It is, therefore, 
frequently overlooked. A further difference is the use of mud for nest building: in 
Cypseloidine swifts it may comprise a considerable proportion (44-89%) of nest 
material (Marin & Stiles 1992), whereas this habit is entirely absent in Waterfall 
Swift (Becking 1971). Finally, there are some osteological differences, particularly 
in respect of the skull (Orr 1963). 

Conclusions 

The resemblance of Waterfall Swift to Cypseloidine swifts is probably due to 
convergence and related to their similar breeding habits. However, the formation of 
a second semiplume-like covering appears to be unique to Waterfall Swift. 
Conversely, differences in nestling plumage between Waterfall Swift and various 
Collocalia and Aerodramus species are probably due to differences in nesting 
habits, rather than large phylogenetic distance. It is unknown what phylogenetic 



Jan-Hendrik Becking 135 Bull. B. O. C. 2006 126(2) 

importance should be attached to the difference in echolocating capabihties between 
Waterfall Swift and various Aerodramus species. DNA studies of H. gigas and 
associated species to date have been inconclusive, and the precise taxonomic 
position of//, gigas remains to be determined. 

Acknowledgements 

I am greatly indebted to Dr Joost Brouwer, Lord Cranbrook and Dr David W. Snow for valuable 
suggestions and comments on an earlier draft. The study would have been impossible without the 
assistance of a semi-permanent crew of Sundanese-Indonesian helpers: Endang, Adi Nurhadi, Maman, 
Odik, Engkus, Adman and Ocang. They were indispensable and should share the outcome of this study. 
I am indebted to them also for their unceasing friendship and care, often under difficult conditions. In 
Sundanese I would say: 'Hatur Nuhun!'. Dr Charles T. Collins, as referee, made many useful comments 
on drafts of this paper. 

References: 

Becking, J.-H. 1971. The breeding of Collocalia gigas. Ibis 113: 330-334. 

Brooke, R. K. 1970. Taxonomic and evolutionary notes on the subfamilies, tribes, genera and subgenera 

of the swifts (Aves: Apodidae). Durban Mus. Novit. 9(2): 13-24. 
Brooke, R. K. 1972. Generic limits in old world Apodidae and Hirundinidae. Bull. Brit. Orn. CI. 92: 

52-57. 
Campbell, B. & Lack, E. 1985. A dictionary of birds. T. & A. D. Poyser, Calton. 
Chandler, A. C. 1916. A study of the structure of feathers, with reference to their taxonomic significance. 

Univ. Calif. Publ. Zool. 13: 243-^46. 
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Address: Ericalaan 7, 6703 EM Wageningen, The Netherlands, e-mail: janhendrikbecking@hotmail.com 
© British Ornithologists' Club 2006 



Donald W. Buden 137 Bull B.O.C. 2006 126(2) 

The birds of Satawan Atoll and the Mortlock 

Islands, Chunk, including the first record of 

Tree Martin Hirundo nigricans in Micronesia 

by Donald W Buden 

Received 1 1 May 2005; final revision received 26 March 2006 

The Mortlock Islands, a subgroup of the Caroline Islands and a part of the Federated 
States of Micronesia (FSM), are among the many small, remote, far-flung and 
seldom-visited islands of Oceania whose faunas are incompletely surveyed; 
ornithological records for the Mortlocks are especially scanty. The present study is 
based largely on observations by the author throughout the Mortlock Islands, but 
mainly on Satawan Atoll, during five separate visits between December 2002 and 
August 2004. 

Previous studies 

Kittlitz (1836) commented briefly on 1 1 species of birds observed on Lukunor Atoll 
in February 1828, during an expedition of the Russian research vessel Senyavin. 
Specimens from this expedition were deposited in the Russian Academy of Sciences 
in St Petersburg (Baker 1951, Hume 2001). Of landbirds recorded on the Mortlocks, 
the St Petersburg collection has only two specimens of Caroline Reed-warbler 
Acrocephalus syrinx that Kittlitz collected on Lukunor (V. Loskot in lift. 2005); 
whether seabirds were included in the collection was not determined at this time. 

Johann Kubary, a naturalist and ethnographer employed at the Godeffroy 
Museum, Hamburg, Germany, visited the southern Mortlocks briefly in 1 873 (Hezel 
1979, Paszkowski 1971), and again in March to late May 1877 (Kubary 1880, 
Schmeltz & Krause 1881). No ornithological reports stem from the 1873 visit, 
which may have been only a brief stop en route from Palau to Pohnpei, but Schmeltz 
& Krause (1881) commented on 19 species seen by Kubary in 1877. Baker (1951) 
included these early records from the Mortlocks in his review of the avifauna of 
Micronesia, but no additional species were added to the group's avifauna by Baker. 

Marshall (1971, 1975) recorded 23 species on Namoluk Atoll based largely on 
observations ancillary to his anthropological studies in 1969-71, which included 
records mentioned by Girschner (1912, 1913), a German physician who conducted 
anthropological research on Namoluk in c.l910 (Marshall 2004). Girschner used 
mainly local vernacular names that Marshall transposed into scientific 
nomenclature. Korte & Meltofte (1997) commented briefly on the status of some 
breeding seabirds in the Mortlocks, but provided no new records. 



Donald W. Buden 



138 



BullB.O.C. 2006 126(2) 



Remarks on place names 

A confusing array of alternative place names for the Mortlock Islands, some 
obsolete, contributes to the difficulty in determining precise locations for some of 
the earlier records. The Mortlock Islands now refers to five atolls and one low 
coralline island south-east of Chunk (=Truk) Lagoon (Fig. 1), but at the time of 
Kubary's visit, in 1877, it referred only to the three southernmost atolls — Ettal, 
Lukunor and Satawan (e.g. Rosser 1870, US Navy Hydrographic Office 1920). The 
name Lukunor is used for one of the islands on the atoll of the same name, and 
Satawan is the name of one of those within Satawan Atoll. Augmenting the 
confusion, Schmeltz & Krause (1881) reported Kubary's records under the heading 
'Die Lukunorgruupe (Mortlock-Inseln)' referring, in this case, to all three closely 
juxtaposed southernmost atolls, and not solely to Lukunor Atoll. Baker (1951), 
however, reported these records as being from Lukunor or Mortlock. But the only 



...... Chuuk 

" . Lagoon 
, ' "j^ : Islands 



MORTLOCK 
ISLANDS 



NAMA 
ISLAND 



I ATOLL 



Chuuk 
Lagoon 



A « 

/ 'y 1 ^ 

^^ Mortlock^ Po 
.0^ 




.10" ^^ v-v ^v 

14tff\Australia 160^ 



NAMOLUK 
ATOLL 



ETTAL 
ATOLL 



50 km 



■■•-^::: '^V* 



SATAWAN J 
ATOLL 

LUKUNOR 
ATOLL 



Figure 1. Location map for the Mortlock Islands. Inset: G = Guam, K = Kosrae, NG = New Guinea, 
Pa = Palau, Po = Pohnpei, Y = Yap. 



Donald W. Buden 139 Bull. B.O.C. 2006 126(2) 

islands specifically mentioned by Schmeltz & Krause (1881) with reference to 
Kubary in the Mortlocks are Ta and Uojta (=Weito), both on Satawan Atoll, and 
Westwood (1905: 81, 112) saw Kubary on Satawan Island in 1877. Also, in an 
anonymously written introduction to Kubary's (1880) ethnographic report on the 
inhabitants of the Mortlock Islands, the following statement appears: 'Kubary 
verweilte auf den Mortlock-Inseln wahrend der Monate Marz biz Ende Mai 1877 
und zwar speciell auf der Inseln Ta, Uoyta und Aliar' [Kubary visited the Mortlock 
Islands, particularly the islands Ta, Weito, and Aliar during the months of March 
through the end of May 1877.]. Additionally, Nason (1970) indicated that Kubary 
spent most of his time on Ta Island during his three months in the Mortlocks. In light 
of these reports, most if not all of Kubary's Mortlock records are likely to have been 
made on Satawan Atoll. 

Study area 

The Mortlocks (05°17'-07°N, 152°35'-153°50'E) comprise a chain of five atolls 
and one low, coral island spanning c.220 km in the west-central Pacific, from Nama 
Island, just south of Chunk Lagoon, south-east to Satawan Atoll (Figs. 1-2). Total 
land area is c. 12 km^ distributed among more than 100 islands; Ta, the largest island, 
is 1.6 km^. Maximum elevations are just c.3-5 m. The Mortlocks lie within the 
equatorial rainbelt and are sufficiently wet to support mesophytic vegetation 
(Mueller-Dombois & Fosberg 1998); the smaller islands lack a freshwater lens and 
are more xeric. Warm humid conditions persist throughout the year. On Namoluk, 
the mean monthly low temperature is 24-2 7''C, and the mean monthly high 30-36°C 
(Marshall 1975). 

Coconut {Cocos nucifera) forest is the dominant vegetation type and breadfruit 
{Artocarpus spp.) is co-dominant in the interior of the larger islands. Other large, 
common forest trees include Barringtonia asiatica, Ficus spp., Guettarda speciosa, 
Hernandia sonora, Neisosperma oppositifolia, Pandanus spp. and Terminalia 
samoensis. The forest abuts the beach or merges abruptly with a narrow zone of 
coastal scrub or thicket dominated by Tournefortia argentea and Scaevola taccada. 
Pemphis acidula scrub is locally common in patches along the shore and mangroves 
are scarce throughout. There is no standing fresh water with the exception of 
excavated wells, some of which are simply crude holes dug to below the level of the 
freshwater lens. Some natural depressions, along with pits excavated for taro 
cultivation, hold water semi-permanently. Large, community-maintained taro 
patches occupy much of the interior on Nama, Namoluk, Kuttu, Moch and 
Lekinioch Islands. 

Methods 

Field work was conducted on Satawan Atoll on 17-26 December 2002, 7 July-1 
August 2003, 30 March-9 April 2004, 22 June-6 July 2004 and 1-5 August 2004. 
The five other groups of islands were visited during summer 2004: Nama Island, 



W. Buden 




140 


Bu 


..#-■::; 


•::;c^;^;^^"-"^'^^-^ 




— j-^S'^SO'N 


.•V." 


'mocH 


'■--■■--. 


153°40'E 


/c^ 




■^* 




:<J— KuttU 






\ . "L Fay Chan 
■ .■ Islands 









Bull.B.O.C. 2006 126(2) 



SATAWAN 
.-.^Onupuku ^^^^^ 



5 km 




Figure 2. Location map for islands on Satawan Atoll, adapted from Bryan (1971). 

7-14 July; Losap Atoll, 10 July; Namoluk Atoll, 19-29 July; Ettal Atoll 30 July-1 
August; and Lukunor Atoll (Lekinioch Island only), 2-3 August. Landbirds were 
counted on 25 islands on Satawan Atoll between December 2002 and August 2004. 
The number of birds seen and heard calling were recorded during timed walks along 
pre-existing trails or by roughly following a compass bearing through the centre of 
the less frequently visited islands, usually along the long axis of the island, but 
occasionally at right angles to it. A single three-minute point count was applied on 
the smallest islands where complete coverage of the island was possible from one 
site. Each transect line or observation point was surveyed only once. An index of 
relative abundance for each species was calculated as the number of individuals 
encountered per hour. Shorebirds and seabirds were counted during walks along the 
beaches at or near low tide. On islands other than Satawan Atoll, birds were 
recorded irregularly and ancillary to reptile and fruit bat surveys; no transect or 
point counts were made, and the species lists, at least of migrants, are incomplete 
for these islands. 

Terms of status and abundance are: common (more than 15 sightings per day), 
fairly common (approximately 5-15 sightings on most days), uncommon (usually 
fewer than five sightings per day and unrecorded on some days), scarce (observed 
occasionally and known only from few records, but not unexpected), resident 
(present year-round and breeding confirmed or very probable), migrant (present 
temporarily or sometimes throughout the year, breeding does not occur or is very 



Donald W. Buden 141 Bull. B.O.C. 2006 126(2) 

unlikely), occasional visitor (non-breeder of irregular and seldom occurrence, but 
not unexpected), vagrant (unexpected visitor and recorded only once or twice), and 
hypothetical (a record in need of corroboration or verification). 

Scientific and English names and sequence of species follow Wiles (2005) 
where applicable. Some authorities include Tree Martin Hirundo nigricans in either 
the genus Petrochelidon or Cecropis. Hypothetical records are in square brackets. 

Species accounts 

[PACIFIC BLACK TiVCV^Anas superciliosa 

Hypothetical. Girschner (1912, 1913) recorded a kind of wild duck [translation from 
German] on Namoluk Atoll. Marshall (1975: 11) stated 'ducks come singly or in 
pairs to the main taro swamp on Namoluk Islet, and that several different people 
reported having seen them at various times during the late 1960's [sicY C. Severance 
{in litt. 2004) also recorded 'ducks' on Losap Atoll during the late 1960s and early 
1970s. In none of these accounts is the species positively identified. Marshall (1971, 
1975) included the Namoluk records under A. poecilorhyncha (= superciliosa) 
probably because of the proximity of Namoluk to the Chunk Lagoon islands, where 
Pacific Black Duck is resident. However, inasmuch as five other species of ducks 
have been recorded as migrants on Chunk and the high islands of Yap and Pohnpei 
that bracket Chunk and the Mortlocks (Wiles 2005), the records of ^. superciliosa 
in the Mortlocks require confirmation.] 

RED JUNGLEFOWL (CHICKEN) Gallus gallus 

Introduced. Free-ranging chickens occur on all the inhabited islands throughout the 
Mortlocks, usually around settlements, but occasionally in more remote areas. Less 
common on some of the uninhabited outlying islands. They were present during the 
earliest surveys. Kittlitz (1836) recorded chickens on Lukunor in 1828 and Kubary 
(in Schmeltz & Krause 1881) noted that Gallus bankiva (=gallus) inhabited the 
interior [probably on Ta or Satawan Islands], where it was rarely disturbed by the 
islanders who used the tail-feathers of roosters as a decoration for combs, but did 
not hunt it for food. In the Mortlocks, chickens are eaten only on special occasions 
and the eggs are very seldom eaten (Tolerton & Rauche 1949?, Marshall 1975, 
Severance 1976). The original sources of introduction are unknown, although 
Tolerton & Rauche (1949?) stated that chickens 'are said to have been brought [to 
Lukunor] from Truk in very early times,' and Marshall (1975) alluded to the 
introduction of Rhode Island Red stock to Namoluk by the Department of 
Agriculture on Truk. 

WEDGE-TAILED SHEARWATER Puffinus pacificus 

Occasional visitor? Encountered widely throughout Micronesia (Wiles 2005) and 
has apparently bred at unspecified localities in the Caroline Islands (Murphy & King 
fide Wiles 2005). Kubary {in Schmeltz & Krause 1881) recorded it in the southern 



Donald W. Buden 142 Bull. B.O.C. 2006 126(2) 

Mortlocks (probably at Satawan Atoll), and Marshall (1971, 1975) remarked on ten 
that arrived at Namoluk Atoll on 17 May 1971 at the same time as a ship from 
Chunk. Marshall (1971, 1975) used the local name 'machukou' for this species, but 
Goodenough & Sugita (1980) indicate the name 'mechikow' (an orthographic 
variant?) may refer to Sooty Tern Sterna fuscata. On the other hand, the name 
'sapal' that Marshall (1975) said referred to an unidentified species of seabird is 
similar to the Chuukese vernacular names 'sepal' (Nomwin Atoll) and 'hepal' 
(PuUap and Puluwat Atolls), which Davis (1999) indicated as possibly referable to 
P. pacificus. 

WHITE-TAILED TROPICBIRD Phaethon lepturus 

Scarce to uncommon resident. Recorded by Kubary {in Schmeltz & Krause 1881), 
presumably on Satawan Atoll, and by Girschner (1912) on Namoluk Atoll. Marshall 
(1975) estimated a population of no more than 50 individuals on Namoluk Atoll and 
recorded them nesting in breadfruit trees ('frequently in clumps of Asplenium 
nidus') on Amwes and Toinom islets. He added that tropicbirds are caught and eaten 
whenever possible. I saw none during this study. 

BROWN BOOBY Sula leucogaster 

Scarce to uncommon, probably breeding in small numbers. Resident breeder and 
widespread in Micronesia (Wiles 2005), but the only one I saw was on Onupuku on 
9 July 2003. Marshall (1975) recorded S. leucogaster as an occasional visitor to 
Namoluk. He used the local vernacular name 'apwang' (='apwan' in Girschner 
1913), but Davis (1999) indicated this is the name for Red-footed Booby Sula sula, 
whereas the name 'aamoo' or 'amo' refers to S. leucogaster. 

RED-FOOTED BOOBY Sula sula 

Locally common resident. I saw Red-footed Boobies frequently among feeding 
flocks of seabirds around Ettal, Satawan and Lukunor Atolls, and estimated 75-100 
breeding pairs (some with eggs and young) on Onupuku Island, Satawan Atoll, on 
9 July 2003. I observed a similar number breeding there on 31 March 2004 when I 
counted 60 active nests, including 29 with downy young, seven with nearly fledged 
young and three occupied by adults. Three immatures kept as pets in the settlement 
on Satawan Island in December 2002 were collected as nestlings on Onupuku. 
Unconfirmed reports of S. sula possibly breeding on Aliar Island, Satawan Atoll 
(Brandt 1962, Nelson 1978, Korte & Meltofte 1997), possibly are referable to 
Onukupu, which is not mentioned in these accounts. The only indications of 
seabirds breeding on Aliar (0.2 ha) that I saw in July 2004 were several disused nests 
of Black Noddies Anous minutus in a large Pisonia tree in the centre of the island. 

GREAT FRIGATEBIRD Fregata minor 

Uncommon to locally common resident. I observed F. minor occasionally 

throughout the year on Satawan Atoll, usually during periods of strong winds and 



Donald W. Buden 143 Bull. B.O.C. 2006 126(2) 

inclement weather, when small flocks of up to 15 soared on obstruction currents 
over the islands. I estimated 25-30 breeding pairs on Onupuku Island on 9 July 
2003: three nests each containing single young ranging in age from a flightless 
downy nestling to fledged or nearly fledged birds with only tufts of down on the 
head and breast, with other (unexamined) nests higher in the canopy beyond reach, 
and some adults flushed from the crowns of coconut trees where additional nests 
may have been hidden from view. I observed a similar number breeding there on 3 1 
March 2004 and recorded a nest with one egg, another with a downy young and five 
others each occupied by an adult. The nests were c.3-6 m above ground in 
Tournefortia trees. Other nests appeared to be in the tops of coconut trees but could 
not be seen clearly and verified. 

INTERMEDIATE EGRET Egretta intermedia 

Occasional visitor? Recorded as a migrant in Micronesia from Palau east to Chunk 

(Wiles 2005). One I saw on Weito, Satawan Atoll, on 28 July 2003 is the first record 

for the Mortlock Islands. It was flushed several times from the marsh (an abandoned 

storm-damaged taro patch) that covers most of the interior of the island. Its larger 

size and slower, more forceful wingbeats distinguished it from the more common 

reef-egret. 

PACIFIC REEF-EGRET Egretta sacra 

Uncommon to fairly common resident. I encountered this species in small numbers, 
of 1-3 birds, regularly on beaches and reef flats throughout the Mortlocks. Of 27 on 
Satawan Atoll for which I recorded coloration, 1 3 were white, six were dark blue 
and eight were piebald. Marshall (1975) estimated no more than 25-30 on Namoluk 
Atoll, and mentioned only the white morph. 

CATTLE EGRET Bubulcus ibis 

Scarce migrant. A migrant to all major island groups of Micronesia (Wiles 2005). 
The two I observed on the Satawan Island school grounds on 17 December 2002 is 
the only record for the Mortlock Islands. 

PACIFIC GOLDEN PLOVER P/wv/a/w/w/va 

Common migrant. I encountered P. fulva regularly on beaches and reef flats 
throughout the Mortlocks, usually in small numbers of up to 5-6 together. They 
were observed throughout the year, but less frequently in summer. 

WANDERING TATTLER Heteroscelus incanus I 

GREY-TAILED TATTLER H. brevipes 

Uncommon to common migrants. Both species are recorded widely in Micronesia 

(Wiles 2005), and because they are difficult to distinguish in the field I treat them 

together. I observed tattlers on beaches throughout the Mortlocks, and usually no 

more than 2-3 together. Two were identified as H. brevipes by their bisyllabic calls; 



Donald W. Buden \AA Bull. B.O.C. 2006 126(2) 

many others had polysyllabic calls more characteristic oiH. incanus. Kittlitz (1836) 
recorded H. brevipes on Lukunor Atoll. Kubary {in Schmeltz & Krause 1881) 
recorded H. incanus, probably on Satawan Atoll, and Marshall (1971, 1975) 
recorded it on Namoluk Atoll. 

COMMON SANDPIPER Actitis hypoleucos 

Uncommon migrant? Kubary {in Schmeltz & Krause 1881) recorded this species in 
the southern Mortlocks, presumably on Satawan Atoll, and Marshall (1971, 1975) 
observed it once on Namoluk Atoll; there are no other records for the Mortlocks. It 
has been recorded widely as a migrant in Micronesia, although Wiles (2005) 
considers records from Pohnpei and Kosrae, in the easternmost Carolines, to require 
confirmation because of the possible occurrence and confusion with the very similar 
Spotted Sandpiper^, macularius (Pyle & Engbring 1985, Wiles 2005). 

WHIMBREL Numenius phaeopus 

Uncommon to fairly common migrant. I observed Whimbrels occasionally on 
Satawan Atoll in April, June, July, August and December, with a max. 5 together at 
Ta airstrip on 4 August 2004. All those flushed at close range clearly had white on 
the rump characteristic of the north-east Asian subspecies A^. p. variegatus. Marshall 
(1971, 1975) recorded this species on Namoluk Atoll. 

BAR-TAILED GOD WIT Limosa lapponica 

Uncommon migrant. Kubary {in Schmeltz & Krause 1881) recorded L. lapponica 

(as L. uropygialis) in the southern Mortlocks, presumably on Satawan Atoll, and I 

saw singles on Ta Island, Satawan, on 7 April and 23 June 2004. There are no other 

records. 

RUDDY TURNSTONE Arenaria interpres 

Common migrant. I frequently encountered Ruddy Turnstones along rocky beaches, 
usually in small flocks of 2-5 (max. 20). It is one of the commonest shorebirds 
during migration and the absence of records from some islands is almost certainly 
an artefact of sampling; recorded previously in the Mortlocks on Namoluk Atoll 
(Marshall 1971) and Lukunor Atoll (Kittlitz (1836). 

BUFF-BREASTED SANDPIPER Tryngites subruficollis 

Vagrant or possibly scarce but regularly occurring migrant. A vagrant in Micronesia 
on Pohnpei and in the Marshall Islands (Wiles 2005). One that I observed foraging 
on the grass airstrip at Ta, Satawan Atoll over several days during the first week of 
April 2004 is the first record for Chunk state. The distinctly buff-coloured plumage, 
yellowish legs and slightly smaller size readily distinguished it from several 
Pluvialis golden plovers foraging in the same area. Its small size and more 
uniformly buffy face and ventral region further distinguished it from Ruff 



Donald W. Buden 145 Bull. B.O.C. 2006 126(2) 

Philomachus pugnax, which is widespread during migration in Micronesia (Wiles 
2005). 

GREAT CRESTED TERN Sterna bergii 

Uncommon to locally common resident. I observed S. bergii on Satawan Atoll 
occasionally, mainly in June and July. One young of the year (with a piece of orange 
yam tied around one leg) on Manimwek Island on 20 July 2003 had a damaged wing 
and was unable to fly. I was told it was captured in the Fauchan Islands group 
(eastern side of the atoll) in March, when there were many flightless young on the 
beach. Kubary {in Schmeltz & Krause 1881) recorded this species, presumably on 
Satawan Atoll, in March-May 1877. Marshall (1975) estimated a population of no 
more than 50 birds on Namoluk Atoll and recorded some of them nesting on 
uninhabited islets. 

BLACK-NAPED TERN Sterna sumatrana 

Uncommon to fairly common resident. I encountered S. sumatrana regularly in 
small numbers throughout Satawan Atoll, mainly on sandy beaches and sandbars. 
Marshall (1975) estimated 50-100 on Namoluk Atoll, where they nested on 
sandbars and exposed reef outcroppings. 

GREY-BACKED TERN Sterna lunata 

Vagrant or occasional visitor. Breeds in Micronesia only in the Mariana Islands, but 
the species has been recorded in Palau, Yap and the Marshall Islands (Wiles 2005). 
One that I photographed (Fig. 3) on a sand and gravel bank on the east side of 
Satawan Atoll on 28 June 2004 is the first record for any of the eastern Caroline 
Islands (i.e. Chunk, Pohnpei and Kosrae states), although Baker (1951) reported that 
W. Coultas obtained an immature male at sea south-east of Pohnpei in October 
1930. The pale grey mantle of the Satawan bird distinguished it from the darker 
plumaged Bridled Tern Sterna anaethetus, which may also occur as a vagrant. 



Figure 3. Grey-backed Tern Sterna lunata, Satawan Atoll, 28 June 2004 (D. W. Buden) 



Donald W. Buden 146 Bull. B.O.C. 2006 126(2) 

BROWN NODDY Anous stolidus 

Common resident. I encountered A. stolidus frequently throughout the Mortlocks. 
Marshall (1975) considered it the most abundant bird on Namoluk Atoll, though I 
found it less numerous than^. minutus. The number of breeders is difficult to assess 
as the nests are usually hidden from view in the tops of coconut and Pandanus trees, 

BLACK NODDY ^«ow5 minutus 

Common resident and probably the most abundant bird in the Mortlocks. I 
frequently observed feeding flocks ranging in size from a few individuals to several 
hundred birds, usually just outside the reef-enclosed lagoons. On Satawan Atoll, in 
July 2003, I estimated c.lOO breeding pairs on Onukupu Island, 200 pairs on Foui 
Island (north-east section of the atoll) and 40 pairs in each of two colonies on Ta 
Island. Smaller colonies of up to c.lO pairs were frequently encountered throughout 
the atoll. Nests usually were placed on the limbs of large broadleaf trees, mainly 
breadfruit and Pisonia grandis. I also frequently observed long, diffuse skeins of 
Black Noddies over Ta airstrip at sunrise, apparently departing their nocturnal roosts 
to feed at sea. I counted roughly 2,115, 2,365 and 2,245 during 05.20-06.40, 
04.45-06.30 and 05.15-06.20 on 24, 25 and 26 June 2004, respectively; sunrise was 
at 05.35. 

WHITE TERN Gygis alba 

Uncommon to locally common resident. I frequently observed White Terns in small 

groups of 2-6 throughout the Mortlocks, usually as they flushed from roosting or 

presumed nesting sites in trees on many of the uninhabited islands and away from 

the villages on the inhabited ones. Marshall (1975) estimated 'well over 700' on 

Namoluk Atoll, and recorded G. alba nesting in breadfruit trees on the three largest 

islets. 

MICRONESIAN PIGEON Ducula oceanica 

Scarce resident, more numerous in the past. The only recent records are from 
Satawan Atoll, where I heard one calling at the eastern end of Ta on 24 December 
2002, and another on Mariong Island on 11 July 2003. Kittlitz (1836) considered i). 
oceanica abundant on Lukunor Atoll, and Kubary {in Schmeltz & Krause, 1881) 
recorded it in the southern Mortlocks, presumably on Satawan Atoll, but gave no 
indication of abundance. Westwood (1905) remarked on shooting at least a dozen 
wild pigeons on an uninhabited island on Lukunor Atoll. Micronesian Pigeon 
probably was extirpated on Lukunor by about the 1940s. Tolerton & Rauche (1949?) 
stated 'some wild pigeons formerly existed but were killed off after acquisition of 
guns by the natives.' Girschner (1912, 1913) indicated D. oceanica was formerly 
present on Namoluk Atoll but extirpated by overhunting. The absence of records 
from Ettal Atoll and of any recent records from Lukunor possibly is an artefact of 
sampling. In view of the widespread distribution of D. oceanica in Micronesia 
(Kosrae, Pohnpei, Chunk, Yap, Palau, Nauru, the Gilbert Islands [= Kiribati] and 



Donald W. Buden 147 Bull. B.O.C. 2006 126(2) 

Marshall Islands; Engbring et al. 1990), indicating an ability to cross long distances 
over water, it seems unlikely that the species would not disperse to Ettal and 
Lukunor from Satawan Atoll across water gaps of less than 10 km. 

POHNPEI LORIKEET Trichoglossus rubiginosus 

Probably introduced but never established; known only from an old record from 
Namoluk Atoll. Girschner {in Marshall 1971) claimed Pohnpei Lorikeet arrived on 
Namoluk during a typhoon in 1905. Marshall (1975) considered it an extinct 
breeder, presumably based on Girschner 's report, but none of the islanders he 
interviewed in 1971 recalled ever seeing this species and there is no evidence it was 
ever established. Pyle & Engbring (1985) suggested the record is more likely based 
on the escape or release of caged birds brought from Pohnpei, where this species is 
endemic and occasionally kept as a pet. 

LONG-TAILED CUCKOO Urodynamys taitensis 

Scarce migrant. Marshall (1975) twice heard this species calling on Namoluk Atoll, 
and Kubary {in Schmeltz & Krause, 1881) recorded it, presumably on Satawan 
Atoll. Long-tailed Cuckoo breeds in New Zealand in the austral summer. It has been 
recorded as a vagrant or migrant in all of the major groups of islands in Micronesia 
(Wiles 2005). 

MICRONESIAN HONEYEATER Myzomela rubratra 

Common resident; the second-most numerous landbird species in the Mortlocks 

after Micronesian Starling Aplonis opaca. Encountered in all terrestrial habitats. 

BARN SWALLOW Hirundo rustica 

Uncommon migrant. Reported during migration on all the major island groups in 
Micronesia (Wiles 2005). I observed H. rustica in small numbers (several groups of 
c.4-6 birds) on Satawan Atoll in December 2000 and April 2004. 

TREE MARTIN Hirundo nigricans nigricans 

Vagrant. A female with small follicles collected at the south-west tip of Satawan 
Island on Satawan Atoll on 1 9 December 2002 is the first record for Micronesia. The 
bird was first observed the previous day hawking back and forth along the beach. It 
was captured in a mist-net set at right angles to the beach. The study skin is 
deposited at the Louisiana State University Museum of Natural Science, Baton 
Rouge (LSUMZ 172489). I saw no other swallows in the vicinity on either day. Tree 
Martin breeds in Australia, Tasmania and two of the Lesser Sundas, and possibly 
southern New Guinea (Coates & Bishop 1997). Migrants (including extralimitals) 
have been recorded as far north as Halmahera in the Moluccas (Buck et al. 1 990) 
and east to New Guinea, the Bismarck Archipelago and the Solomons (Peckover & 
Filewood 1976), and New Caledonia (Barre & Bachy 2003). The wing-chord 
(unflattened, left 109.8 mm, right 109.1 mm) of the Satawan specimen falls well 



Donald W. Buden 148 Bull B.O.C. 2006 126(2) 

within the range of the nominate (eastern Australian) subspecies (105-111 mm — 
Schodde & Mason 1999), but outside those of the western Australian subspecies H. 
n. neglecta (100-107 mm — Schodde & Mason 1999), and the Lesser Sunda Islands 
race H. n. timoriensis (90-94 mm — Turner & Rose 1989). The December date of the 
Satawan record is unusual for an austral migrant as it falls within the austral summer 
breeding season. 

[UNIDENTIFIED HIRUNDINIDAE 

Small groups of 5-10 swallows that I observed in flight on Ta and Satawan Islands 
in December 2002 and April 2004 were unidentified to species. They lacked white 
on the rump, thus eliminating Tree Martin. In having a slightly forked tail and 
seemingly no dark pigmentation on the upper breast they appeared less similar to 
Bam Swallow than to Pacific Swallow H. tahitica, which occurs in south-east Asia 
south to Australia, New Zealand and Melanesia, and east to the Society Islands in 
Polynesia (Howard & Moore 1994), but is unrecorded in Micronesia.] 

CAROLINE REED-WARBLER Acrocephalus syrinx 

Common resident. Recorded throughout the Mortlocks and frequently encountered 
in dense understorey and thickets. Marshall (1975) estimated 400-500 on Namoluk 
Atoll. Kubary {in Schmeltz & Krause 1881) recorded it nesting in March-May 
1 877, presumably on Satawan Atoll. I saw two nearly fledged young that a boy had 
removed from a nest on Ta Island, Satawan Atoll, on 7 July 2003. 

MICRONESIAN STARUNG Aplonis opaca 

Common resident; the most common landbird on Satawan Atoll (Table 1), Namoluk 
Atoll (Marshall 1975), and elsewhere throughout the Mortlocks (pers. obs.). 
Marshall (1975) recorded it nesting on Namoluk Atoll, usually in Pandanus and 
coconut trees. On Satawan Atoll, I saw at least six different immatures in striped 
plumage soliciting food from adults in June 2004, and an islander informed me of 
the discovery of two unfledged young in a nest in the crown of a coconut tree on 
Manimwek Island on 18 July 2003. 

[BLUE-FACED PARROTFINCH Erythrum trichroa 

Hypothetical. Baker (1951) questionably recorded this species from Lukunor 
probably based on a statement by Schmeltz & Krause (1881) relating to Kubary 's 
work in the Mortlocks: 'Ein versch lagenes Exemplar der Erythrura trichroa schuss 
K. einmal hier [Kubary once shot a specimen oi Erythrura trichroa here].' But the 
location of any such specimen is unknown. E. trichroa occurs on the high Caroline 
Islands, including Palau, Chunk, Pohnpei and Kosrae, but it is otherwise unknown 
from the atolls. The record from the Mortlocks (stated as Lukunor but probably 
Satawan) seems unlikely and requires confirmation.] 



Donald W. Buden 



149 



Bull.B.O.C. 2006 126(2) 



TABLE 1 
Distribution of birds in the Mortlock Islands' 





Nama 


Losap 


Namoluk 


Ettal 


Lukunor 


Satawan 


Species 


Island 


Atoll 


Atoll 


Atoll 


Atoll 


Atoll 


[Anas superciliosa] 






G/M 








Gallus gallus^ 


+ 


+ 


+/G/M 


+ 


+/K 


+/SK 


Puffinus pacificus* 






M 






SK 


Phaethon leptunis * 






G/M 






SK 


Sula leucogaster* 






G(?)/M 




K(?) 


+ 


Sula sula* 






G(?) 






+ 


Fregata minor* 






G/M 






+/SK 


Egretta intermedia 












+ 


Egretta sacra* 




+ 


+/G/M 


+ 


+/K 


+/SK 


Bubulcus ibis 












+ 


Pluvialis fulva 


+ 


+ 


+/G/M 


+ 


+/K 


+ 


Heteroscelus incanus 


+(?) 


+(?) 


G(?)/M(?) 






+/SK 


Heteroscelus brevipes 


+(?) 


+(?) 


G(?)/M(?) 




K 


+ 


Actitis hypoleucos 






M 






SK 


Numenius phaeopus 






G/M 






+/SK 


Limosa lapponica 












+/SK 


Arenaria interpres 


+ 


+ 


+/G/M 


+ 


K 


+ 


Tryngites subruficollis 












+ 


Sterna bergii* 






G/M 






+/SK 


Sterna sumatrana* 






G/M 






+/SK 


Sterna lunata 












+ 


Anous stolidus* 


+ 


+ 


+/G/M 


+ 


+/K 


+/SK 


Anous minutus* 


+ 


+ 


+/G/M 


+ 


+ 


+/SK 


Gygis alba* 






G/M 




K 


+/SK 


Ducula oceanica* 






G 




K 


+/SK 


Trichoglossus rubiginosus^' 






G 








Urodynamys taitensis 






M 






SK 


Myzomela rubratra* 


+ 


+ 


+/G/M 


+ 


+ 


+ 


Hirundo rustica 












+ 


Hirundo nigricans 












+ 


Acrocephalus syrinx* 


+ 


+ 


+/G/M 


+ 


+/K 


+/SK 


Aplonis opaca* 


+ 


+ 


+/G/M 


+ 


+/K 


+/SK 


[Erythrura trichroa] 












SK 



Sources: + = this study, G = Girschner {in Marshall 1975), K = Kittlitz (1836), M = Marshall (1975), 

SK = Kubary {in Schmeltz & Krause,1881). 

Symbols and abbreviations: [] = hypothetical record; * = indigenous species recorded breeding in the Mortlocks 

or on one or more islands elsewhere in the Carolines; ' = introduced. 



Discussion 

Of the 33 species of birds recorded from the Mortlock Islands (Table 1), and 
excluding the two introduced species and two hypothetical records, 15 breed locally 



Donald W. Buden 




150 


Bull. B.O.C. 2006 126(2) 






TABLE 2 






Relative abundance of landbird 


species on Satawan Atoll based on surveys on 


25 islands. 








Encounter rate (birds/h)'' 






Inhabited 


Uninhabited 




Species 


Incidence^ 


islands' 


islands'^ X^' 


All islands 


Ducula oceanica 


1/25 (4%) 





0.2 (1) NA 


0.1 


Myzomela rubratra 


25/25 (100%) 


14.1 (49) 


17.2(75) ns 


15.8 


Acrocephalus syrinx 


24/25 (96%) 


9.5 (33) 


14.6 (64) ns 


12.4 


Aplonis opaca 


24/25 (96%) 


22.4 (78) 


26.1(114) ns 


24.5 



' The number of islands on which the species was recorded, divided by the total number of islands surveyed. 

'' The number of birds seen or heard per hour; total number in parentheses — see Methods. 

" Four: Moch, 0.28 km-, 50 minutes; Satawan, 1.1 km", 60 minutes; Ta, 1.6 km^, 40 minutes; Kuttu, 0.28 km\ 60 

minutes. 
•^ Twenty-one: total area c.1.1 km- (largest island 0.13 km-), total survey time 262 minutes (4.37 hours). 
'•■ Comparisons between inhabited and uninhabited islands: NA = not available, insufficient data, ns = no significant 

difference (P >0.05). 

or have been recorded breeding elsewhere in the Caroline Islands; none is endemic. 
The 14 others are regularly occurring migrants or occasional visitors and vagrants. 
The indigenous avifauna includes 11 seabirds in the following orders: 
Procellariiformes (one shearwater), Pelecaniformes (two boobies, a frigatebird and 
a tropicbird), and Charadriiformes (six terns). All but one of the terns {Sterna 
lunata) breeds on one or more islands in the Carolines. The eight species of 
shorebirds recorded in the Mortlocks are all boreal migrants, and, with three egrets 
and a duck, complete the list of water or marsh birds. In view of the relatively large 
number of migrant shorebirds recorded on the major island groups elsewhere in the 
FSM (29 species — Wiles 2005), the number recorded in the Mortlocks is likely to 
increase markedly with additional surveys. 

Onupuku, a small (0.05 km^) island isolated on the west side of Satawan Atoll 
(Fig. 2), is an important seabird rookery in the Mortlocks. Satawan islanders 
consider it the principal bird island on the atoll. Onupuku is the chief breeding site 
for Red-footed Boobies in the Mortlocks (75-100 pairs); Great Frigatebirds, Brown 
Noddies, Black Noddies and White Terns also nest there. Onupuku is part of the 
Kuttu Island district and permission to visit must be obtained from a family on 
Kuttu. Some poaching of birds and coconut crabs occurs, but for the most part 
islanders are respectful of the restrictions, which are based on cultural tradition 
rather than formal Western legislation. 

The four resident landbirds represent three feeding guilds, including a nearly 
extirpated frugivore, Micronesian Pigeon, apparently now confined to a small 
population on Satawan Atoll, another frugivore (Micronesian Starling), an 
insectivore (Caroline Reed-warbler) and a nectarivore (Micronesian Honeyeater). 
The honeyeater, reed-warbler and starling are abundant and widespread throughout 
the Mortlocks, and their encounter rates on Satawan Atoll showed no significant 



Donald W. Buden 151 Bull. B.O.C. 2006 126(2) 

differences between the larger, inhabited islands and the smaller, uninhabited ones 
(Table 2). The six other landbirds include introduced chickens, a probably 
introduced but apparently never established lorikeet, two migrants (Long-tailed 
Cuckoo and Bam Swallow), a vagrant (Tree Martin) and one hypothetical (Blue- 
faced Parrotfmch). Additional surveys will doubtless increase the number of 
migrants and occasional visitors, but the four resident landbirds undoubtedly are all 
that occur. This same resident species complex, or subsets thereof, is typically 
present on atolls throughout the eastern Caroline Islands (Baker 1951, Buden 1995, 
1996a,b, 1998, 1999a,b), the one exception being Micronesian Kingfisher 
Todiramphus cinnamominus on Ant Atoll, off Pohnpei (Engbring et al. 1990, Buden 
1996a). 

Acknowledgements 

I thank Jeremiah Trimble for confirming identification of the Sterna liinata, Waher Boles, Steve Cardiff 
and Ron Clouse for assistance with the literature, S. Cardiff for providing measurements of the 
subspecies of Hinindo nigricans, and Vladimir Loskot for information on Mortlock Islands specimens at 
the Russian Academy of Sciences. 

References: 

Baker, R. H. 1951. The avifauna of Micronesia, its origin, evolution, and distribution. Univ. Kans. Publ. 

Mus. Nat. Hist. 3: 1-359. 
Barre, N. & Bachy, R 2003. Complement to the list of the birds of New Caledonia. Alauda 71: 31-39. 
Bryan, E. H. 1971. Guide to place names in the Trust Territory of the Pacific Islands. Pacific Science 

Information Center, Bishop Museum, Honolulu. 
Buck, H., King, B. & Yong, D. 1990. Tree Martin Hirundo nigricans sighting on Halmahera. Kukila 5: 

76. 
Buden, D. W. 1995. Reptiles, birds, and mammals of Mokil, and Pingelap atolls, eastern Caroline Islands. 

Micronesica 28: 9-23. 
Buden, D. W. 1996a. Reptiles, birds, and mammals of Ant Atoll, eastern Caroline Islands. Micronesica 

29: 21-36. 
Buden, D. W. 1996b. Reptiles, birds, and mammals of Pakin Atoll, eastern Caroline Islands. Micronesica 

29: 37^8. 
Buden, D. W. 1998. The birds of Kapingamarangi Atoll, including first record of the Shining Cuckoo 

{Chrysococcyx lucidus) from Micronesia. Notornis 45: 141-153. 
Buden, D. W. 1999a. Reptiles, birds, and mammals of Oroluk Atoll, eastern Caroline Islands. 

Micronesica 31: 289-300. 
Buden, D. W. 1999b. The birds of Sapwuahfik Atoll, with first record of the Grey Wagtail Motacilla 

cinerea from the Federated States of Micronesia. Bull. Brit. Orn. CI. 119: 261-270. 
Coates, B. J. & Bishop, K. D. 1997. The guide to the birds ofWallacea. Dove Publications, Alderley. 
Davis, A. E. 1999. A preliminary list of animal names in the Chuuk District, with some notes on plant 

names. Micronesica 32: 1-245. 
Engbring, J., Ramsey, F. L. & Wildman, V. J. 1990. Micronesian forest bird surveys, the Federated States 

of Micronesia: Pohnpei, Kosrae, Chuuk, and Yap. US Fish & Wildlife Service, Honolulu. 
Girschner, M. 1912. Die Karolineninsul Namoluk und ihre Bewohner. Baessler-Archiv 2: 123-215. 
Girschner, M. 1913. Die Karolineninsul Namoluk und ihre Bewohner. Baessler-Archiv 3: 165-190. 
Goodenough, W. H. & Sugita, H. 1980. Truk-English dictionary. American Philosophical Society, 

Philadelphia. 
Howard, R. & Moore, A. 1994. A complete checklist of the birds of the world. Second edn. Academic 

Press, London. 



Donald W. Buden 152 Bull. B.O.C. 2006 126(2) 

Hume, J. P. 2001. Notes on the extinct Kosrae Starling ^/?/o«« corvina Kittlitz, 1833. Bull. Brit. Orn. CI. 

122: 141-154. 
Kittlitz, F. H. 1836 [reprinted 1971]. Observations Zoologiques faites pendant I'expedition de la corvette 

Le Seniavine. Pp. 237-330 in Lutke, F. (ed.) Voyage autour du monde, 1826-1829, vol. 3. 

Bibliotheca Australiana No. 60. Da Capo Press, New York. 
Korte, J. de & Meltofte, H. 1997. Notes on breeding sites of Pelecaniformes in Micronesia. Bird Conserv. 

Intern. 7: 287-290. 
Kubary, J. 1880. Die Bewohner der Mortlock-Inseln. Mitt. Geog. Gesellschaft Hamburg (1878-1879): 

224-299. 
Marshall, M. 1971. Notes on birds from Namoluk Atoll. Micronesica 7: 234-236. 
Marshall, M. 1975. The natural history of Namoluk Atoll, Eastern Caroline Islands. Atoll Res. Bull. 189: 

1-53. 
Marshall, M. 2004. Namoluk beyond the reef. Westview Press, Boulder, CO. 
Paszkowski, L. 1971. John Stanislaw Kubary^ — ^naturalist and ethnographer of the Pacific islands. 

Austral. Zool. 16: 43-70. 
Peckover, W. S. & Filewood, L. W. C. 1976. Birds of New Guinea and tropical Australia. A. H. & A. W. 

Reed. Sydney. 
Pyle, P. & Engbring, J. 1985. Checklist of the birds of Micronesia. Elepaio 47: 11-15. 
Rosser, W. H. 1870. The seaman 's guide to the islands of the North Pacific. North Pacific Pilot, part 2. 

James Imray & Son, London. 
Schmeltz, J. D. E. & Krause, R. 1881. Ein Beitrag zur Kunde der Siidsee-Volker. Die Ethnographisch- 

Anthropologische Abtheilung des Museum Godeffroy in Hamburg. L. Friederichsen & Co., 

Hamburg. 
Schodde, R. & Mason, I. J. 1999. The directory of Australian birds: passerines. CSIRO Publishing, 

CoUingwood, Victoria. 
Severance, C. J. 1976. Land, food, and fish: strategy and transaction on a Micronesian atoll. Ph.D. thesis. 

Dept. of Anthropology, Univ. of Oregon, Eugene. 
Tolerton, B. & Rauch, J. 1949? Social organization, land tenure, and subsistence economy ofLukunor, 

Nomoi Islands. Coordinated Investigation of Micronesian Anthropology (CIMA) Report No. 26. 

Office of Naval Research & The National Academy of Sciences, Washington DC. 
Turner, A. & Rose, C. 1989. A handbook to the swallows and martins of the world. Christopher Helm, 

London. 
US Navy Hydrographic Office. 1920. Pacific islands pilot, vol. 1. Second edn. Government Printing 

Office, Washington DC. 
Westwood, J. 1905. Island stories; being extracts from the papers of Mr John Westwood, mariner, of 

London and Shanghai. North-China Herald Office, Shanghai. 
Wiles. G. J. 2005. A checklist of the birds and mammals of Micronesia. Micronesica 38: 141-189. 

Address: Division of Natural Sciences and Mathematics, College of Micronesia — FSM, P.O. Box 159, 
Kolonia, Pohnpei 96941, Micronesia, e-mail: don_buden@comfsm.fm 

© British Ornithologists' Club 2006 



Diego F. Cisnews-Heredia 153 Bull. B.O.C. 2006 126(2) 

Notes on breeding, behaviour and distribution of 
some birds in Ecuador 

by Diego E Cisneros-Heredia 

Received 28 May 2005; final revision received 29 March 2006 

The avifauna of Ecuador is reasonably well known in terms of distribution and 
relative abundance (Ridgely & Greenfield 2001). Geographic patterns are rather 
well established (e.g. Fjeldsa & Krabbe 1990, Ridgely & Greenfield 2001), but 
breeding biology data for most Neotropical species is poor, and available 
information general and incomplete. Quantitative and qualitative understanding of 
breeding biology and ecology is essential for the development of efforts to stop the 
ongoing loss of biodiversity. I present novel data on the breeding biology and 
distribution of 48 taxa, including several species considered rare or 
globally/nationally threatened by Ridgely & Greenfield (2001), Granizo et al. 
(2002) or BirdLife International (2004). Whilst my data are incomplete, the 
usefulness of such natural history observations is considerably augmented when 
combined with those of others, permitting us to establish country-wide patterns of 
reproductive biology and seasonality (Greeney et al. 2004). 

Material and methods 

Information presented herein was obtained between 1990 and 2005 at various 
localities in the Pacific lowlands, Andean highlands and Amazonian lowlands of 
Ecuador (Table 1, Fig. 1). Coordinates and elevations were confirmed using the 
physical map published by the Institute Geografico Militar in 2000. Taxonomy, 
English names and distributional information follow Remsen et al. (2006) and 
Ridgely & Greenfield (2001). I use the following abbreviations: (B) building, (I) 
incubating, (N) nestlings, (F) dependent fledglings, (AN) active nest at unknown 
stage, (J) juvenile-plumaged individual and (CM) carrying nesting material but nest 
unseen. 

Species accounts 

COMMON PIPING-GUAN Aburria pipile 

I observed an adult with a chick under its wing, from a canopy tower c.30 m above 
ground, at Tiputini Biodiversity Station, in December 1997. Hilty & Brown (1986) 
reported females from Colombia in breeding condition in February and laying birds 
in May. My observation involved the taxon cumanensis, considered specifically, as 
Blue-throated Piping-guan Aburria (or Pipile) cumanensis, by Sibley & Monroe 
(1990), Rodner et al. (2000) and Grau et al. (2005). 



Dieso F. Cisneros-Heredia 



154 



Bull. B.O.C. 2006 126(2) 



TABLE 1 
Localities mentioned in the text (see Fig. 1). 



Loc# 


Localities, provinces 


Coordinates 


Altitude (m) 


1 


Comuna Nueva Juventud, Sucumbios 


c.00°05'S, 76°12'W 


C.290 


2 


Cuenca-Loja road, near Saraguro, Loja 


c.03°36'S,79°13'W 


c.2,600 


3 


Cuicocha Lagoon, Imbabura 


00°18'N,78°22'W 


3,100 


4 


Cumbaya, Pichincha 


00°12'S, 78''25'W 


2,390 


5 


Jipijapa-Puerto Lopez road, Manabi 


c.0r24'S, 80°45'W 





6 


La Joya (Hacienda), Pichincha 


00°05'N, 78°59'W 


750-800 


7 


La Merced de Nono (Hacienda), Pichincha 


c.00°02'S, 78°33'W 


3,100-2,750 


8 


La Perla Protected Forest, Esmeraldas 


00°0rN, 79°22'W 


150 


9 


Malacatos, Loja 


04°14'S, 79°17'W 


c. 1,500 


10 


Mashpi Protected Forest, Pichincha 


00°09'N, 78°50'W 


1,050 


11 


Rio Guajalito Protected Forest, Pichincha 


00°14'S, 78°49'W 


1,900 


12 


Rio Palenque Scientific Center, Los Rios 


00°35'S,79°21'W 


200 


13 


San Antonio de Pichincha-Perucho road, Pichincha 


c.00°02'N, 78°25'W 


c.2,200 


14 


Tandayapa, Pichincha 


c.00°0rS, 78°46'W 


c. 1,700 


15 


Tinalandia, Pichincha 


00''18'S,79°03'W 


740 


16 


Tiputini Biodiversity Station/ River, Orellana 


00°39'S, 76°08'W 


190-250 


17 


Vilcabamba, Loja 


04°13'S,79°15'W 


1,600 


18 


Yuturi Lodge, Orellana 


00°33'S, 76°05'W 


200 


19 


Yanacocha, Pichincha 


00°10'S, 78°35'W 


3,400 


20 


Papallacta, Napo 


00°22'S, 78°13'W 


3,600 


21 


Quito (including Bosque Protector Pichincha and 








Parque Metropolitano de Quito), Pichincha 


00°irS,78°29'W 


2,800-3,000 


22 


La Toreadora Lagoon, Parque Nacional Cajas, Azuay 


c.02°52'S, 79°13'W 


3,600 


23 


La Cienega (Hacienda), Cotopaxi 


00°45'S, 78°40'W 


3,000 


24 


Bosque Protector Pasochoa 


00°27'S, 78°29'W 


2,900 



DARK-BACKED WOOD-QUAIL Odontophorus melanonotus 
A pair with at least two dependent chicks at Mashpi Protected Forest ( 1 8 km north 
of San Miguel de Los Bancos) on 9 March 2003. When approached the adults fled 
and began calling vigorously, whilst the chicks quickly took cover under roots or 
amidst leaf litter, and froze. They were in secondary forest with a dense, tangled 
understorey and fairly open canopy on steep slopes. In March 2002, a pair with at 
least three dependent chicks was observed in a road cut covered by dense 
vegetation, c.l5 km south (by road) from Rio Guajalito Protected Forest. When 
approached, the entire group hid amidst vegetation. Strewe (2001) noted downy 
chicks in June 1997 and December 1997, and young in September 1997, January 
1997 and January 1998 at La Planada, Colombia, and Greeney & Nunnery (2006) 
recorded juveniles in May from the Tandayapa area. These observations suggest 
year-round breeding in north-west Ecuador and south-west Colombia. O. 
melanonotus is classed as Vulnerable in Ecuador and globally (Granizo et al. 2002, 
BirdLife International 2004). 



Diego F. Cisneros-Heredia 



155 



Bull. B.O.C. 2006 126(2) 



Pacific 
Ocean 



Colombia 



8* 



^5^ 



.12. 



.^ 



• 10 
24 



23 



i13 



20 



• 1 



•^18 



16 



22 



Pert/ 



17 



100 km 




Figure 1. Map of Ecuador showing the study sites; numbers correspond to the localities in Table 1, except 
for Quito (locality 21) which is represented by a rectangle between localities 4 and 19 for clarity. 



PIED-BILLED GREBE Podilymbus podiceps 

At least six adults at Cuicocha Lagoon (00°18'N, 78°22'W; 3,100 m) on 17 
November 2001, including a pair with downy young. Most birds were c.30-50 m 
from the dock of the lagoon, but the pair with chicks was near shore. The species 
breeds year-round: in Colombia, nests have been reported in January-March, egg- 
laying peaks in January-March and September-October, and pairs with downy 
chicks have been recorded in July- August (Hilty & Brown 1986, Fjeldsa & Krabbe 
1990). 



Diego F. Cisnews-Heredia 1 56 Bull. B. O. C. 2006 126(2) 

MAGNIFICENT FRIGATEBIRD Fregata magnificens 

In August 1995 and June 2000, birds were observed in courtship activity in the 
crowns of trees along the road between Jipijapa and Puerto Lopez. Haase (1997) 
reported that the courtship season in Ecuador usually commences in September; my 
records thus precede the normal breeding season by up to three months. However, 
in September 2005 no evidence of such activity was observed. Haase (1997) 
suggested that the early start in July 1997 of this species' courtship was influenced 
by the 1997 El Niiio Southern Oscillation (ENSO), but my observations were made 
in non-El Nino years. Although variations on the courtship season of i^ magnificens 
could be related to ENSO or to 'La Nina' (eastern tropical Pacific Ocean cooling), 
more data are needed to assess direct and indirect effects on the species' breeding 
biology. 

AMERICAN KESTREL Falco sparverius 

A resident pair at Cumbaya, Universidad San Francisco de Quito campus, 
constructed a nest within dense ivy on the wall of a building, c.l2 m above ground. 
The pair was first observed frequently entering the nest in late July 2004. Chicks 
were heard in early September. Both sexes were apparently active at all stages of 
nesting, but nestling provision was by the male alone whilst the female always 
guarded the nest (similar observations have been reported for F. rufigularis by Sick 
1993). Several times there were confrontations with Great Thrushes Turdus fuscater 
which attempted to enter the nest. Both sexes engaged in nest defence, but the male 
was more aggressive and followed the thrushes up to 75 m. The species nests on 
rock-ledges or in cavities, such as holes in trees, posts, cliffs, embankments or 
termite nests (Hilty & Brown 1986, Fjeldsa & Krabbe 1990, Sick 1993). The ivy had 
a depth of c. 15-20 cm and the nest was in a cavity formed by the foliage. Breeding 
in Ecuador has been reported in June and July (Fjeldsa & Krabbe 1990). 

ANDEAN / SLATE-COLOURED COOT Fulica ardesiaca 
At least three nests under construction and another with one egg amongst floating 
vegetation at Cuicocha Lagoon on 17 November 2001. Breeding has been reported 
in late February in Nariiio, Colombia (Hilty & Brown 1986). Fjeldsa & Krabbe 
(1990) mentioned that egg-laying peaks in July-August, but that clutches may be 
initiated year-round. 

BARN OWL Tyto alba 

A fledgling at Hacienda La Cienega, on 25-26 June 2005, amidst the leaves of a 
large palm tree, c.6 m above ground, and attended by one adult at night. Several 
egragopiles that were collected contained mainly hair and bones of Mus musculus, 
Rattus rattus and an unidentified rodent species. Ridgely & Greenfield (2001) 
reported T. alba as recorded mainly below 2,000 m, and that higher records perhaps 
correspond to wandering birds, citing just one record, from Hacienda La Cienega. 
At least two T. alba used to live at Parque La Carolina, Quito, at 2,800 m. They were 



Diego F. Cisneros-Heredia 157 Bull. B.O.C. 2006 126(2) 

apparently resident, being noted monthly in 1995-2004. A T. alba was periodically 
recorded at Parque Ingles, Quito, at c.3,000 m, and at least two periodically heard 
near Avenida Occidental, Quito, in 1994-2000. At least one T. alba was regularly 
recorded at Quebrada Ashintaco, Parque Metropolitano de Quito. These records 
indicate some highland populations of T. alba are resident as high as 3,000 m. 

SAND-COLOURED NIGHTHAWK Chordeiles rupestris 
An unsexed adult incubating two eggs at Tiputini Biodiversity Station, August 2000. 
The nest was a simple depression in the sand, c.5 m from the bank of the rio 
Tiputini, almost beneath a broken log. The eggs resembled those described by Hilty 
& Brown (1986). Nests on sand bars have been reported in July-late September in 
the Leticia area of Colombia (Hilty & Brown 1986). 

WHITE-TIPPED SWIFT Aeronautes montivagus 

I found a colony beside the San Antonio de Pichincha-Perucho road on 1 October 
1994, close to where Marin (1993) observed the species. The colony comprised c.40 
birds and was located in a vertical road cut (c.50 m high). Several flying juveniles 
were seen at the entrances to nests, confirming breeding. 

STRIPE-THROATED HERMIT Phaethornis striigularis 

An adult feeding a juvenile in a forest gap with low ferns at Mashpi Protected 
Forest, 18 km north of San Miguel de Los Bancos, on 8 March 2003. The juvenile 
had the rump and back entirely rufous, unlike the adult, which has the back bronzy- 
green and the rump rufous. 

LITTLE WOODSTAR Chaetocercus bombus 

A pair observed at Tandayapa Lodge on 20 August 2003. The female was perched 
on an outer branch of an Inga tree whilst the male performed a courtship display 
consisting of perpendicular flights in front of the female together with high-pitched 
calls and wing-sounds. The maximum height of the flight above the female was c.5 
m. This behaviour was observed for almost 20 minutes. 

AMAZONIAN WHITE-TAILED TROGON Trogon viridis 
An adult female excavating a hole in the lower part of an arboreal termite nest, c.40 
m above ground, at Tiputini Biodiversity Station in July 1999. Due to the position 
of the nest entrance, I assume the nest was burrowed as depicted by Sick (1993: 
353), i.e. the bird creates an upward-sloping tunnel to an incubation chamber in the 
centre of the nest. A male in breeding condition has been recorded in June in 
Colombia (Hilty & Brown 1986). 

RED-HEADED BARBET Eubucco bourcierii 

An adult male was carrying food to a cavity c.2.5 m high in a 5-m dead stump, in a 
gap within secondary forest, at Rio Palenque Scientific Center, on 10 November 
2004. Nestling calls were heard from within. A male Black-cheeked Woodpecker 



Diego F. Cisneros-Heredia 



158 



Bull. B.O.C. 2006 126(2) 




Figure 2. Palm with nest (lower right) of Cranioleiica cmtisiensis, Mashpi Protected Forest, Pichincha, 
Ecuador, 8 March 2003. The black arrow points to the leaf to which the nest was attached (Diego F. 
Cisneros-Heredia) 



Diego F. Cisneros-Heredia 



159 



BullB.O.C. 2006 126(2) 




Figure 3. Cross-section of nest of Cranioleuca antisiensis, Mashpi Protected Forest, Pichincha, Ecuador, 
8 March 2003 (Diego F. Cisneros-Heredia). A = Roof of brood chamber (top). White bar indicates nest 
entrance (bottom). The roof of the brood chamber is covered by brown hairy Bombacaceae seeds (lower 
photo). 



Diego F. Cisneros-Heredia 160 Bull. B.O.C. 2006 126(2) 

Melanerpes pucherani was observed inspecting the same hole, cA hour prior to this 
observation, but no interactions were observed. 

LINE-CHEEKED SPINETAIL Cranioleuca antisiensis 

A pair of Cranioleuca carrying material to a nest at Mashpi Protected Forest, 1 8 km 
north of San Miguel de Los Bancos, on 8-9 February 2003. The nest was a bulky 
structure attached to the backside tip of a lower, outermost leaf of a palm, c. 1 m 
above ground. The palm (c.l5 m high) was surrounded by bushes on a hillside 
covered in foothill evergreen forest (Fig. 2). Following afternoon rain on 9 February 
2003, both birds were carrying moss and palm fibres to the nest (repairing?). On 8 
March 2003, the nest was abandoned and contained several fragments of broken 
eggshell (predation?). The nest coincides with Zyskowski & Prum's (1999) 
description for a pensile nest. It was a domed structure externally constructed of 
moss and strips of a cucurbitaceous vine, with palm fibres and dry Lauraceae leaves 
incorporated within the interior. The inner chamber was oval, wider than tall (height 
10 cm, width 15 cm). The nest entrance was an 8-cm downward-sloping tunnel at 
the bottom of the nest. The brood chamber was lined with rootlets of an epiphyte 
(Ericaceae) and vines, and the dome covered with hairy seeds of a Bombacaceae 
(Fig. 3). The plumage of the nesting birds was typical of C. antisiensis, which in 
Ecuador is known only from the south-west: rufous crown, white supercilium and 
whitish throat, which characters differentiate it from juvenile Red-faced Spinetail C. 
erythrops. Previous reports from north-west Ecuador are regarded as uncertain due 
to the resemblance of C. antisiensis to juvenile C. erythrops (Ridgely & Greenfield 
2001). 

GOLDEN-FACED TYRANNULET Zimmerius chrysops I 
YELLOW-BROWED TODY-FLYCATCHER Todirostrum chrysocrotaphum 
A nest of each species was observed, from a canopy tower, in an emergent Ceiba 
sp., C.45 m above ground, at Tiputini Biodiversity Station in August 2000. The nests 
were just c.l m apart and c. 1 m from a paper wasp (Vespidae) nest. Only one adult 
was seen at the nest of T. chrysocrotaphum, but both at the nest of Z chrysops. No 
interactions were observed between the two species. Similar observations for nests 
of T. chrysocrotaphum located near a wasp nest were reported by Greeney et al. 
(2004). 

CHESTNUT-COLLARED SWALLOW Petrochelidon rufocollaris 

A large colony (c.300 birds) nesting on a church in Malacatos on 31 December 

2002. Numerous nests, at various stages of construction, were observed and 

photographed. Several birds were observed flying over Vilcabamba the same day. 

The species was not previously recorded from the Malacatos area and its presence 

supports the suggestion by Ridgely & Greenfield (2001) that the species is 

increasing. 



Diego F. Cisneros-Heredia 161 Bull. B.O.C. 2006 126(2) 

WHITE-CAPPED DIPPER Cinclus leucocephalus 

An active nest in a hole within a rock- wall dam of the rio Guajalito, in late 
July-early August 1998, at Rio Guajalito Protected Forest. Both sexes entered the 
nest several times per day, presumably feeding the single chick. The pair used the 
same nest location in two consecutive years in the same months, but in 1999 two 
chicks fledged. The pair disappeared in 200 1 , probably due to a petroleum spill and 
the release of hundreds of trout by the owner of a commercial open aquaculture, and 
there were no further observation until at least late 2003. 

CASQUED OROPENDOLA Clypicterus oseryi 

A colony of c.30 pairs was discovered high (c.SO m) in an emergent tree beside the 
rio Tiputini, at Tiputini Biodiversity Station, in January 1998. In January 1999 the 
colony was smaller (c.l5 pairs), but active, and remained so at least until January 
2000. The colony was checked in the months of January-March, June- August, and 
December, and breeding activity noted in March, August and December. 

LESSER GOLDFINCH Carduelis psaltria 

A flock of 11 at Vilcabamba, on 30 December 2003, was photographed as they 
moved through the canopy of low Acacia trees and shrubs in a lodge garden near a 
ravine. The flock comprised adults of both sexes, and two dependent juveniles 
whose plumage corresponded to the description in Fjeldsa & Krabbe (1990). In 
Colombia, birds in breeding condition have been reported in May-October, eggs in 
May, July, December and January, fledglings in March, May and August, and 
juveniles in April, May and September (Hilty & Brown 1986, Fjeldsa & Krabbe 
1990). C. psaltria is very local and usually rare in the foothills and subtropical zone 
on both slopes of the Andes, with only one recent report from Pichincha province 
(Ridgely & Greenfield 2001). I have the following observations; six at La Merced 
de Nono, on the Nono-Calacali road, on 1 October 1994; three near Tandayapa on 
13 November 1994; and a pair at the Rio Guajalito Protected Forest in August 1998. 
All observations involved birds foraging in the canopy of trees or tall shrubs. 

ORANGE-BELLIED EUPHONIA Euphonia xanthogaster 
A pair attending nestlings at Hacienda La Joya on 9-10 January 2002. The nest was 
a globular mossy ball (c.l5 cm diameter) with a side entrance, as previously 
described for the species (Hilty & Brown 1986), at the end of a lone branch c.1.5 m 
above a fast-moving rivulet (c.6 m wide). Both adults were active dawn to dusk. The 
male never entered the nest, but was always close by (c.50 cm), occasionally 
vocalising. The female constantly returned at intervals of 0.25-2.0 hours, and 
remained within the nest for c. 10-1 5 -minute periods. The adults exhibited a 
complex behaviour when the female approached (observed more than 1 5 times on 
two consecutive days). The female arrived, always on branches higher or lower than 
the nest, and flew between different branches, from back to front and left to right. 



Diego F. Cisnews-Heredia 162 Bull. B.O.C. 2006 126(2) 

for c.\ minute, before pausing at a branch posterior to the nest, whilst the male 
always perched over the nest. The female swiftly flew to the right, to the nest, whilst 
the male simultaneously performed an elliptical flight, intersecting with the female 
just in front of the nest, before returning to its perch. These movements seemed to 
represent anti-predator behaviour, using distracting movements when the female 
arrived and confounding it with male flight (a hypothesis also presented by Sargent 
1993, who suggested that a mate-guarding component might be involved). The 
female was never seen carrying food, but both adults were mist-netted and the 
female regurgitated fruits and seeds, probably mistletoe berries. My observations 
were made with C. Dingle and G. Castaiieda. Such behaviour has been described for 
several other Euphonia (Isler & Isler 1987, Sargent 1993). H. F. Greeney {in litt. 
2005) has also noted similar behaviour in this species, and for Ochraceous Attila 
Attila torridus (Greeney 2006). E. xanthogaster in breeding condition have been 
reported in February-July and nests in November-April in Colombia (Hilty & 
Brown 1986). 

Other records include: Andean Guan Penelope montagnii, 29 June 1996, 2 F, 
La Merced de Nono (with two adults in a forest remnant adjacent to cattle pasture). 
Rufescent Tiger-heron Tigrisoma lineatum, 17 July 2000, 1 J, Comuna Nueva 
Juventud (captured at night in a net placed for bats). Andean Condor Vultur 
gryphus, 23 July 1995, 1 J, Papallacta (with an adult male). White Hawk 
Leucopternis albicollis, 15 August 1996, 1 AN, Yuturi Lodge (large stick platform 
nest in crown of an emergent Ceiba sp. with one adult in attendance). Eared Dove 
Zenaida auriculata, 28 August 2005, 1 AN and 2 F on AN, Centro Cultural 
Metropolitano de Quito (females attending both nests on a palm tree leaf c. 10 m 
above ground). 2 November 2005, 1 AN, Jardin Botanico de Quito (nest c.3 m 
above ground). Blue-and-yellow Macaw Ara ararauna, 16 August 1996, 1 AN, 
Yuturi Lodge (on a tall palm stump beside a river). Hoatzin Opisthocomus hoazin. 
May 2001, 1 N and 1 AN, Tiputini Biodiversity Station (beside an oxbow lake). 
Great Potoo Nyctibius grandis, 27 December 1999, 1 AN, Tiputini Biodiversity 
Station (nest in crown of an emergent Ceiba sp. beside the rio Tiputini). Rufous 
Potoo A^. bracteatus, 28 December 1999, 1 N, Tiputini Biodiversity Station (day 
roost and nest on stump in forest gap c.8 m above ground in lowland evergreen 
seasonally flooded forest). Golden-headed Quetzal Pharomachrus auriceps, 21 
August 1999, 1 AN, Rio Guajalito Protected Forest (female carrying food to hole in 
trunk c.lO m above ground). Pacific Hornero Furnarius cinnamomeus, 29 June 
1996, 1 AN, Hacienda La Joya (nest c.25 m above ground). Blackish Tapaculo 
Scytalopus latrans, 20 August 1 990, 1 J, Bosque Protector Pasochoa. Tufted Tit- 
tyrant A nairetes parulus, 9 August 1994, 1 J, Bosque Protector del Pichincha, Quito 
(mist-netted with two adults, juvenile had yellow rictus). Tumbesian Tyrannulet 
Phaeomyias tumbezana, 4 October 2004, 1 F, Universidad San Francisco de Quito, 
Cumbaya campus (in bushes beside artificial lagoon). White-tailed Shrike-tyrant 
Agriornis andicola, 28 December 2002, CM, Cuenca-Loja road, near Saraguro, 



Diego F. Cisneros-Heredia 163 Bull B.O.C. 2006 126(2) 

Loja (in cattle pasture surrounded by low bushes, pine and eucalyptus). Masked 
Water-tyrant Fluvicola nengeta, July 1998, 1 AN, La Perla Protected Forest (adult 
attending globular nest, 5 m above ground in a 3-m tall bush in a field adjacent to 
cattle pasture). Black-crowned Tityra Titym inquisitor, CM (7 January 1995), AN 
(until April 1995), Hacienda La Joya (pair with nest in a hole atop a c.l5 m stump 
in cattle pasture less than 10 m from a forest fragment). White-thighed Swallow 
Neochelidon tibialis, 8 March 2003, 3 F, Mashpi Protected Forest (flock of c.l5 with 
two Southern Rough- winged Swallow Stelgidopteryx ruficollis). Great Thrush 
Turdus fuscater, 2 November 2005, 2 CM, Termas de Papallacta. Hooded 
Mountain-tanager Buthraupis montana, 22 June 2005, 2 F, Yanacocha (in 
subcanopy and understorey as part of a mixed flock). Cinereous Conebill 
Conirostrum cinereum, 30 November 2005, 1 CM, Jardin Botanico de Quito. Tit- 
like Dacnis Xenodacnis pahna, 15 August 1999, F and J, La Toreadora Lagoon 
(with adults in PolylepislGynoxys forest). Glossy Flowerpiercer Diglossa 
lafresnayii, 22 June 2005, 2 F and 1 J, Yanacocha (with adults at hummingbird 
feeders). Saffron Finch Sicalis flaveola, 30 December 2003, AN, Vilcabamba (pair 
occupying abandoned Furnarius cinnamomeus nest, c.8 m above ground). Yellow- 
breasted / Rufous-naped Brush-finch Atlapetes latinuchus, 20 August 1990, 1 I, 
Bosque Protector Pasochoa. Russet-crowned Warbler Basileuterus coronatus, 20 
August 1990, 1 AN, Bosque Protector Pasochoa. Hooded Siskin Carduelis 
magellanica, 8 August 1994, 2 J, Bosque Protector del Pichincha, Quito (mist-netted 
with an adult female, juveniles had red rictus). 

Acknowledgments 

I thank Harold Greeney and Jesse Delia for their helpful comments on a draft of this paper. I am 
extremely grateful to Maria Elena Heredia and Laura Heredia for their constant, loving support (moral, 
financial, logistical) of my work during the past ten years; to Kristof Zyskowski, Francisco Veintimilla 
and Mercedes Foster for access to literature; the British Ornithologists' Club for my free membership in 
2004, enabling access to valuable literature. I acknowledge staff of the Cotacachi-Cayapas Ecological 
Reserve, Aves & Conservacion (Corporacion Omitologica del Ecuador), and Fundacion Natura for their 
support on the 2001 Cuicocha trip, and 1995 and 2000 visits to Machalilla National Park. Thanks to 
Marcelo Negrete and Susana Izurieta for granting access to Hacienda La Joya; Tiputini Biodiversity 
Station, especially David & Consuelo Romo, for continuous financial, logisdc and moral support; and 
Universidad San Francisco de Quito for institutional and logistical support. For field companionship, I 
express my gratitude to Maria Elena & Laura Heredia, Andres Baquero, Freddy Caceres, Roberto 
Phillips, Mayer Rodriguez, Chris Schneider, Charlie Kieswetter, Caroline Dingle, Gabriela Castaneda, 
Jeffrey Arellano, and my partners and professors of the College of Biological & Environmental Sciences, 
Universidad San Francisco de Quito (1997-2004), especially Margarita Brandt, Susana Cardenas, 
Heman Castaneda, Angel Chiriboga, Karina Dammer, Andres Leon, Pablo Melo, Diego Mosquera, 
Karina Newman, Gunther Reck, Javier Robayo, Geovanna Robayo, David Romo, Vlastimil Zak and Toa 
Loaiza. I thank Paul Greenfield and Niels Krabbe for their patience, support and mentorship over the 
years. 

References: 

BirdLife International. 2004. Threatened birds of the world 2004. CD-ROM. BirdLife International, 
Cambridge, UK. 



Diego F. Cisnews-Heredia 164 Bull. B.O.C. 2006 126(2) 

Fjeldsa, J. & Krabbe, N. 1990. Birds of the high Andes. Zool. Mus., Univ. of Copenhagen & Apollo 

Books, Svendborg. 
Granizo, T., Pachecho, C, Ribadeneira, M. B., Guerrero, M. & Suarez, L. (eds.) 2002. Libro Rojo de las 

aves del Ecuador, vol. 2. SIMBIOE/Conservacion Internacional/EcoCiencia/Ministerio del 

Ambiente/UICN, Quito. 
Grau, E. T., Pereira, S. L., Silveira, L. R, Hofling, E. & Wajntala, A. 2005. Molecular phylogenetics and 

biogeography of Neotropical piping guans (Aves: Galliformes): Pipile Bonaparte, 1856 is a synonym 

of Aburria Reichenbach, 1853. Mol. Phyl. & Evol. 35: 637-645. 
Greeney, H. F. 2006. The nest and eggs of the Ochraceous Attilay^mVa torridus in south-west Ecuador 

with notes on parental care. Cotinga 25: 60-62. 
Greeney, H. F., Gelis, R. A. & White, R. 2004. Notes on breeding birds from an Ecuadorian lowland for- 
est. Bull. Brit. Orn. CI. 124: 28-37. 
Greeney, H. F. & Nunnery, T. 2006. Notes on the breeding of north-west Ecuadorian birds. Bull. Brit. 

Orn. a. 126: 38^5. 
Haase, B. 1997. The impact of the El Nino Southern Oscillation (ENSO) on birds: update from Ecuador 

1997. Cotinga 8: 64-65. 
Hilty, S. L. & Brown, W. L. 1986. A guide to the birds of Colombia. Princeton Univ. Press. 
Marin, M. 1993. Patterns of distribution of swifts in the Andes of Ecuador. Avocetta 17: 117-123. 
Remsen, J. V., Jaramillo, A., Nores, M., Pacheco, J. F., Robbins, M. B., Schulenberg, T. S., Stiles, F. G, 

Silva, J. M. C, Stotz, D. F. & Zimmer, K. J. 2006. A classification of the bird species of South 

America. http://www.museum.lsu.edu/~Remsen/SACCBaseline.html. 
Ridgely, R. S. & Greenfield, R J. 2001. The birds of Ecuador, vol. 1. Cornell Univ. Press, Ithaca, NY. 
Rodner, C, Lentino, M. & Restall, R. 2000. Checklist of the birds of northern South America. Pica Press, 

Robertsbridge. 
Sargent, S. 1993. Nesting biology of the Yellow-throated Euphonia: large clutch size in a Neotropical fru- 

givore. Wilson Bull. 105: 285-300. 
Sibley, C. G & Monroe, B. L. 1990. Distribution and taxonomy of birds of the world. Yale Univ. Press, 

New Haven. 
Sick, H. 1993. Birds in Brazil: a natural history. Princeton Univ. Press. 

Skutch, A. F. 1996. Antbirds and ovenbirds, their lives and homes. Univ. of Texas Press, Austin. 
Zyskowski, K. J. & Prum, R. O. 1999. Phylogenetic analysis of the nest architecture of Neotropical oven- 
birds (Fumariidae). Auk 116: 891-911. 

Address: Colegio de Ciencias Biologicas y Ambientales, Universidad San Francisco de Quito, Casilla 
Postal 17-12-841, Quito, Ecuador, & Aves & Conservacion (Corporacion Omitologica del Ecuador), 
e-mail: diegofrancisco_cisneros@yahoo.com 

© British Ornithologists' Club 2006 



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COMMITTEE 

Cdr. M. B. Casement OBE, RN {Chairman 2005) I. R. Bishop OBE (2003) 

Miss H. Baker (Vice-Chairman) (2005) C. W. R. Storey (2003) 

S. A. H. Statham {Hon. Secretary) (2004) J. R Hume (2004) 

D. J. Montier {Hon. Treasurer) (1997) Dr J. H. Cooper (2005) 

R J. Wilkinson (2005) 
Ex-officio members 

Hon. Editor. G. M. Kirwan (1 January 2004) 

Chairman of BOU-BOC Joint Publications Committee (JPC): Revd. T. W. Gladwin 
Hon. Website Manager. S. R Dudley (2005) 
BOC Publications Officer Prof R. A. Cheke (200 1 ) 
Chairman of the Bulletin Subcommittee: Prof C. J. Feare (2005) 



Bulletin of the British Ornithologists' Club 

ISSN 0007-1595 
Edited by Guy M. Kirwan 

Volume 126, Number 2, pages 81-164 

CONTENTS 

Club Announcements 81 

DONEGAN. T. M. & HUERTAS, B. A new brush-fmch in the Atlapetes latinuchus complex from 

the Yariguies Mountains and adjacent Eastern Andes of Colombia 94 

BECKING, J.-H. Notes on Waterfall Swift Hydrochous gigas: I. Occurrence and 

nesting 117 

BECKING, J.-H. Notes on Waterfall Swift Hydrochous gigas: II. Nestling plumage 

and phylogenetic relationships 123 

BUDEN, D. W. The birds of Satawan Atoll and the Mortlock Islands, Chuuk, including the first 

record of Tree Martin Hirundo nigricans in Micronesia 137 

CISNEROS-HEREDIA, D. F. Notes on breeding, behaviour and distribution of some birds in 

Ecuador 153 



CONTENTS 

Authors are invited to submit papers on topics relating to the broad themes of taxonomy and distribution of 
birds. Descriptions of new species of ^irds are especially welcome and will be given priority to ensure rapid 
publication, subject to successful passage through the normal peer review procedure, and they may be 
accompanied by colour photographs or paintings. On submission, manuscripts, double-spaced and with 
wide margins, should be sent to the Editor, Guy Kirwan, preferably by e-mail, to GMKirwan@aol.com. 
Alternatively, two copies of manuscripts, typed on one side of the paper, may be submitted to the Editor, 74 
Waddington Street, Norwich NR2 4JS, UK. Where appropriate half-tone photographs may be included and, 
where essential to illustrate important points, the Editor will consider the inclusion of colour figures (if pos- 
sible, authors should obtain funding to support the inclusion of such colour illustrations). As far as possible, 
review, return of manuscripts for revision and subsequent stages of the publication process will be undertak- 
en electronically. For instructions on style, see the inside rear cover of Bulletin 125 (1) or the BOC website 
www.boc-online.org 

Registered Charity No. 279583 

©British Ornithologists' Club 2006 

www.boc-online.org 



Printed on acid-free paper. 

Pubhshed by the British Ornithologists' Club 

Typeset by Alcedo Publishing of Pennsylvania, USA, and printed by Crowes of Norwich, UK